the vegetation of rock fissures, screes, and … · 2.materials and methods 2.1. study area the...

STUDIA GEOBOTANICA 10: 15-58, 1990

THE VEGETATION OF ROCK FISSURES, SCREES, AND SNOW-BEDS IN THE PIRIN PLANINA MOUNTAINS (BULGARIA)

LadislavMUCINA,MilanVALACHOVIC,lvanJAROLIMEK,JanSEFFER,AnnaKUBINSKA & Ivan PISUT

Keywords: alpine vegetation, Bulgaria, endemie plants, syntaxonomy, Aspleni,etea trichomanis, Salicetea· herbaceae, Thlaspietea rotundifolii

Abstract: The vegetation of calcareous rock fissures and crevices, marble screes, and snow-beds on granite, marble and lime-rich schists was studied in the Pirin Planina Mts., SW Bulgaria. Three plant communities of the rock-fissure vegetation were described, including the Hi-eracio pannosi-Caricetum, Leontopodio-Potentilletum stojanovii, and the Silene pusilla-Saxifraga oppositifolia community (both in subalpine and alpine altitudes). The syntaxonomic relations of the communities to the Ramondion nathaliae were discussed. The scree vegetation on marble in the alpine belt is represented by the Papaveri-Armeri,etum (with severa! subassociations and variants) which populates shady and moist habitats, and the Veronico kellereri-Silenetum prostratae growing on sunny, dry, south-facing screes. They were classified within a new alliance, the Veronico-Papaverion degenii, an endemie unit for Bulgaria. The low-altitude screes were classified within the Bromo lacmonices-Geranietum macrorrhizi (Silenion marginatae). Snow-bed vegetation on silicate bedrock belongs to the Salicion herbaceae. The Ligustico - Plantaginetum gentianoidis (concave relief forms,long-lasting snow cover) and the Omalotheco -Alopecuretum gerardii (convex reìief forms, short-lasting snow cover ), were described from this alliance. The marble snow-beds in the Pirin Planina Mts. are classified within the Salicion retusae which includes two associations such as the Bartsio-Salicetum reticulatae and Gentiano-Plantaginetum atratae,characteristic of the long-lasting and short-lasting snow patches,respectively. Ali of the associations, butthe Leontopodio-Potentilletum stojanovii and Ligustico-Plantaginetum gentianoidis, were described as new. Ali of the communities house many species endemie to the Pirin Planina Mts ., Bulgaria and Balkan Peninsula.

1. Introduction

The high-mountain vegetation of the Balkan Peninsula is an amazingly variedsystem of "cold islands in the sea of warm air". It is assumed that during the Pleistocene the psychrophilous and oligothermic vegetation in the Balkan Peninsula covered larger areas, and these might have been interconnected also with the Alps and Carpathians. Although the Continental Glacier was not reaching as far as the Balkans, there were many local glaciers covering particular mountain ranges of the Rodopi-Dinarid Region (Frenzel 1967).

After the retreat of the glaciers to higher altitudes and their decay, the territory covered by psychrophilous vegetation shrinked and "an archipelago of the cold islands in the sky" emerged. These geologica! and climatica! processes gave push to allopatric speciation in these small areas and many endemie plants were formed or

15

old relict species were saved (Walter & Straka 1970). The "archipelago" is an excellent example of evolutionary laboratory well

suited for studies of vicariance-biogeography aspects as well as vegetationchorologic phenomena (Horvat et al. 1974, Krahulec 1985). As the alpine and subalpine vegetation of the Balkans is better preserved than that of the Alps, the Carpathians or the Scandinavian mountains, it is also more diversified. This is also reflected in abundance of endemie alliances, orders, and in Greece even classes (Quézel 1964).

The syntaxonomy of the high-mountain vegetation in the Balkans was performed especially in those countries where the Braun-Blanquet approach of floristicsociological classification of vegetation set up stable roots. These countries include also Yugoslavia where Horvat and many of his pupils and colleagues elaborated the mountain vegetation in numerous papers (for references see Horvat et al. 1974). Much syntaxonomic work has been done in Rumania in the high-mountain areas (Domin 1933, Borza 1959, Pu�caru-Soroceanu et al. 1963, 1981, Boi:ìcaiu 1971, Sanda et al. 1977, Schneider-Binder & Voik 1977, Resmerita &Pop 1984, Coldea 1984, 1985, volume of Comunicari de Botanica, Bucurei:ìti, 1977, to mention some important ones). There are a few syntaxonomical papers on the high-mountain vegetation also from Greece (Quézel 1964, 1967). In Bulgaria, on the other hand, vegetation scientists were using other methods of vegetation classification (see for instance Bondev 1959, 1966, Ganchev 1963, Juhasz-Nagy 1963, Kochev 1967, Kochev & Nikolov 1976 etc.).

The Pirin Planina Mts. is touristically one of the most popular high-mountain resorts of Bulgaria (the Vikhren Mt. is considered the National Mountain). A national park was declared to preserve the nature of this unique area, and this park was included into the list of Biosphere Reserves (Goodier & Jeffers 1981). In 1958 a seminal paper by Simon was published on the alpine vegetation of the Pirin Planina Mts. This paper remained, however, the only reliable published information on floristic-sociological syntaxonomy of the mountains till today.

Our paper is aimed to supply data on other alpine and subalpine vegetation types of the Pirin Planina Mts. than those given by Simon (1958), and to present a syntaxonomical discussion on the described vegetation types. Some ideas on synecology of the types were presented elsewhere (l\,1ucina et al. 1986).

2. Materials and Methods

2. 1. Study Area

The Pirin Planina Mts. is situated in the southwestern Bulgaria. It is limited by the valley of the Struma (Strimon) River from the west, by the Predel Saddle from the north, by the Mesta River from the east and by the Parilska Sedlovina Saddle from Slavyanka Mts. It is the third highest mountain range of the Balkan Peninsula. The highest peak reaches 2915 m (Vikhren Mt. or El-Tepe Mt.). The total area of the range is 1210 km2 (Dushkov 1983). Pirin Planina Mts. is classified into three

16

parts such as Northern, Central, and Southern. The northern part is situated between the Predel Saddle (1140 m) and the Todorova Polyana Saddle (1883 m), while the central part is divided from the southern one by the Popovi Livadi Saddle (1400 m).

The main mountain range follows the direction NW to SE and ramificates into

numerous side-ridges. The NW slopes are very steep, the SE slopes are gentle. The altitudinal belts in the mountains are represented as follows: 600 to 1000 m above sea level - 25%, l000to 1600 - 3 8%, 1600to 2200 - 24%,and altitudes above 2200 -

13 %. Three of the peaks reach above the alti tu de 2900 m (Vikhren, Kutelo I - 2907 m

and Kutelo II - 2908 m).

The mountain range is a part of the Rila -Rodo pi Massif built of crystalline rocks belonging to an old Thracic-Macedonian Massif, and sedimentary rocks (mainly carbona tic) of the Mesosoic ori gin. The high-mountain modelation of the relief is a

result of the glacial activity of a local glacier in the Pleistocene. The main range of the Pirin Planina Mts. in its northern part is built of granitoid rocks, crystalline

schists and marbles. The main area of the latter is situated between the Kabata Saddle ( 2700 m) and Kamenititsa Saddle (app. 2600 m). The marbles are grey or grey-white, and stratified by dark strips. Due to long-term tectonic activities they are disintegrated and broken (Georgiev 1956).

The climate is typical of high-mountain altitudes as the most of the area (over 75%) is situated at altitudes above 1000m. The highest altitudes are characterized by low temperature, small yearly temperature amplitude, windiness (the prevailing winds are from W and S W directions), high air humidity, long-lasting snow cover, and intensive irradiation. The average yearly temperature in Sandanski ( 275 m) is 13.3 ° C, in Bansko ( 963 m) 9.0° C. On the contrary, the temperature of the coldest month (January) is - 4.2°C (measured at the Vikhren Cottage). For a comparison, the J anuary temperature in Sandanski (in the Struma V alley) is 3 .4° C. The warmest month is August ( 8. 7° C at the Khizha Vikhren (Cottage) and 23.5° C in Sandanski). The total precipitation in the area is much higher than that in the adjacent basins. The Sandanski Basin is characterized by yearly precipitation of 541 mm, the Razloshki Basin by 702 mm, while 1571 mm was ascertained at the Vikhren Cottage. The cloudeness is the highest in May (around 70%). Snow cover lasts from 5 to 6 months. There is a permanent snow field considered the only Balkan glacier in the Golemiya Kazan Circle. The snow cover is very uneven (Mironski et al. 1970).

The rivers, brooks and rills emerging in the Pirin Planina Mts. belong to two river basin. The southwestern slopes are drained by the Struma River while the northeastern slopes are drained by the Mesta River. The largest tributaries of the Struma include the Pirinska Bistritsa, Sandanska Bistritsa, Vlakhinska Reka and Melnishka Reka. The Mesta River collects waters of the By ala Reka, Glazne, Retize and Tufcha. The maximal point water stands of the rivers falls within the month of May when the snow fields above timberline are thawing (Mironski et al. 1970).

The Pirin Planina M ts. is well-known for the variety of glacial lakes counting around 17 6 in summer.

17

The studi ed area belongs phytogeographically to the Central European Floristic Region, Balcanic Subregion, Province of Moesiacum orientale, region of Pirin

(Bondev et al. 1973). Our phytosociological material comes from the northern part of the area (the

crystalline limestone and marble ridge) and some valleys and ridges of the central part (Fig. 1).

2.2. Field Work

The field work was done during 3 phytosociological expeditions to southBulgarian mountains (Pirin Planina and Rila Planina Mts .) in July to August in 1978, 1983 and 1984. The relevés were taken from physiognomically relatively homogeneous stands found in homogeneous habitats. The area of the relevés was

adjusted to the extent of well-developed stands and kept approximately the same for the same vegetation type.

By sampling seven-grade scale ofBraun-Blanquet (1964) ranging from rto 5was used in 1978, while the grade 2 was subdivided into 2 m, 2 a and 2 b in sampling the alpine vegetation in 1983 and 1984 (according to Barkman et al. 1964).

2.3. Data Treatment

The critical plant taxa were determined according to Flora na Bulgariya (Stoyanov & Stefanov 1948), Flora na NR Bulgariya (Yordanov 1963-1979) and Flora Europaea by Tutin et al. (1964-1980). The nomenclature of phanerogams follows generally Flora Europaea (see Halliday & Beadle 1983 for a list). The nomenclature of mosses follows Frahm & Frey (1983), and that of lichens is according to Wirth (1980).

The abundance- cover classes recorded in the field were unified by using a

transformation of van der Maarel (1979) into a scale ranging from O to 9. The transformed relevés were entered into community phytosociological tables.

Traditional syntaxonomic methods of the Braun -Blanquet approach, as re vis ed by Westhoff &van der Maarel (1978), were used in the synthetic phase of the work. We adopted character species, differential species and the diagnostic species

combination for the characterization and delimitation of the syntaxa, where by also characteristics of habitats were partly taken into consideration as secondary classification criteria.

English romanization was used in transliteration of original Bulgarian names written in cyrilics.

3. Plant Communities

Three groups of plant communities were investigated in the present paper, such as (1) communities of rock rissures of the Asplenietea trichomanis, (2) communities

. 18

PLANINA f91RIN MTS.

Hm

I. o

e u

. thern Bulgaria.

. Mts. m sou p· · Planma

. an e system of the mn f the mam r g F. 1-A mapo

9

1g.

1

of marble screes of the Thlaspietea rotundifolii, and (3) communities of snow beds on siliceous as well as lime-rich bedrocks of the Salicetea herbaceae.

3.1. Plant Communities in Rock Fissures and Crevices

In the following paragraphs we bring description and syntaxonomic discussion on plant communities in rock fissures and crevices classified within the Asplenietea

trichomanis. Only the communities of lime-rich (marble and crystalline limestone schists) in the northern range on the Pirin Planina Mts. were studied. Three communities were distinguished such as the Hieracio pannosi-Caricetum kitaibelianae Gower altitudes), Leontopodio-Potentilletum stojanovii (higher altitudes), and the Silene pusilla-Saxifraga appositi/olia community (schists).

Hieracio pannosi-Caricetum kitaibelianae ass. nova Nomenclatura! type: Tab. 1, relevé 7, hoc loco

The community was recorded at lower altitudes (subalpine and oreal belts) in the close surroundings of the Banderitsa and Vikhren Cottages.

The slopes, rocky ridges and stone walls above the Banderitsa Cottage are formed by crystalline limestones and marbles. At lower altitudes, at around 1900 and 2000 m, these are overshadowed by a Pinus leucodermis wood. In the krummholz belt they become exposed to direct insolation and strong winds. The slope aspects and inclination reflect in species distribution pattern. Gentle slopes and broken stony crests are mainly covered by a grassland formation dominated by Sesleria coerulans and Carex kitaibeliana. Species -rich stands are supported by steep rock faces where the plants dwell in rock crevices and fissures, especially in half-shaded habitats. Asplenium ruta-muraria, A. trichomanes, Kernera saxatilis, Campanula cochleariifolia var .pirinica Vel. and Saxifraga paniculata are typical in these habitats. The open stands of the communities are penetrated by Sesleria coerulans, Hypochoeris pelivanovicii Petr .,Minuartia verna, Helianthemum nummu

larium subsp. tomentosum, Euphrasia salisburgensis and Paronychia kapela. The Hieracio pannosi-Caricetum kitaibelianae is limited to eastern aspects of

the steep slopes and crests which are rich in fissures and small rocky terraces. The character taxa of the community include Hieracium pannosum, Asperula pirinica Stoj. et Acht., Globularia meridionalis, Draba lasiocarpa, Allium flavum var. minus

Boiss. and Centaurea mannagettae (I'ab. 1, relevés 1-9). The moss cover is very small Gess than 5%); only Tortella tortuosa occurs more

frequently.

Leontopodio-Potentilletum stojanovii Simon 1958 Nomenclatura! type: Simon (1958: Tabelle I, relevé 1), lectotypus

This association replaces the Hieracio pannosi-Caricetum kitaibelianae at altitudes above 2 300 m. The typical habitats of the community are insolated marble

20

outcrops and steep, prevailingly south-facing, slopes. Potentilla apennina subsp. stojanovii is the dominant taxon of the community.

This chamaephytic dwarf shrub forms compact carpets in some places covering also the bare rocks. It is characterized by a stout and branched woody rhizone which as a worm penetrates the fissures and crevices in marble.

The distribution area of Potentilla apennina is limited to mountains of the central Apennines (eg. Majella Mts .) and those of the Balkan Peninsula (see below). Flora Europaea (Ball et al. 1968) distinguishes two subspecies ,P. apennina subsp. a pennina (Italy) and subsp. stojanovii (Bulgaria and Greece ). The Saxifrago-Potentilletum apennincie was reported from The Sar, Jakupica and Nidza Mts. (Horvat 1936, see also Horvat et al. 1974). Jovanoviè - Dunjiè (1953-1955) described the Potentillo apenninae - Saxifragetum paniculatae from the Suva Mts. However, the species cannot be considered characteristic of the latter association as it penetrates also other Ramondion nathaliae communities, eg .Erysimo -Ramondietum nathaliae in that region (Tab. 2). Frequent companions are Campanula cochleariifolia var. pirinica, Saxifraga juniperifolia, Leontopodium alpinum subsp. nivale, Thymus cherlerioides and Helianthemum canum (Tab. 1, relevés 10-16). Many FestucoSeslerietea species are also found in stands of the Leontopodio-Potentilletum stojanovii. Unlike in the Hieracio pannosi-Caricetum kitaibelianae, more cryptogams are found in the Leontopodio-Potentilletum stojanovii, such as Tortella tortuosa, Hymenostylium recurvirostre, Toninia candida. The average cover of mosses and lichens is 5-10%.

Silene pusilla -Saxifraga oppositifolia community

In the area of the Bayuvi Dupki (2820 m) and the Kamenititsa Mt. (2710 m) a rare community Silene pusilla - Saxifraga oppositifolia is encountered. This community is limited to steep wall outcrops composed of parallel strata of lime-rich schists. The cracks separating the schist blocks are containing plenty of water, thus the bare rock-face (usually from 0.5 to 2 m high) resembles a low-capacity waterspring as the rock-face is soaked with water even when the surrounding habitats are dry.

Several Saxifraga (sub) species are found to share the site: Saxifraga appositi/olia, S. adscendens subsp. discolor ry el.) Kuzm., S. luteoviridis, S. exarata subsp .pirinica(S. Pawl.) Kuzm. and S. ferdinandi-wburgi (Tab. 3). All of them have theircoenologic optima in other communities (screes, alpine grasslands), but this isprobably the only safe site they can grow together. The fissures house smallspecimens of Galium stojanovii, Asplenium fissum and Silene pusilla which, when inoptimal phenological stage, dictate the appearance of the community. The strikingspecies-richness of mosses and lichens is also characteristic of the community.Among other species, the Thlaspietea rotundifolii diagnostic taxa, such as Poapirinica, Veronica saturejoides subsp. kellereri (see Fischer & Fischer 1981),Papauer degenii (Urum. et Jav .) Kuzm., Thlaspi bellidifolium, Arabis caucasica andDoronicum columnae are frequently found here.

21

Tab. 1 - Plant communities of the Asplenietea trichomanis in the Pirin Planina Mts.

No. of relevés 1111111

123456789 0123456

Exposition E s E

NNS s s s s

EEEEEEEEE WSESEEE

Slope 0 755686784 8578786

000500005 0000000

Sarnpled area m2 1 1121 2

969272585 9012293

Cover of herb layer % 656546434 2361634

000000000 5005000

Cover of moss layer % <<<< 1 « 1<

111151055 5515505

HIERACIO-CARICETUM KITAIBELIANAE

Hieracium pannosum (Ch) Hypochoeris pelivanovicii {Ch) Asperula suberosa (Ch) Carurn riqidulum agg. (Ch) Centaurea mannagettae (Ch) Ailium * minus (Ch) Globularia cordifolia (Ch) Micromeria cristata (Ch) Minuartia verna (D) Teucrium montanum (D) Helianthemum * tomentosum (Dl Paronychia kapela (D) Sedum album (D) Dianthus petraeus (D) Knautia sp. {D) Draba lasiocarpa (D) Stipa pulcherrima (Dl

422423432 ..... 12

222323332 ..... 2.

222.23322 ..... 22

2.233.222 ..... 2.

2222.32.2 ...... 2

2222.22 ..

.. 3.53535

22224.2 ..

3322.4232 ...... 2

2243 ... 35

2342 .. 22 . . 2 .....

32 ... 2.22

2.23.35 ..

2. 2 .. 25 ..

. 2. 2 .. 2. 2

2 .. 2 .. 2 .. .... 25 .. 3

LEONTOPODIO-POTENTILLETUM STOJANOVII

Potentilla * stojanovii (Chl Saxifraga juniperifolia IChl Hymenostylium recurvirostre (O) Fulgensia bracteata (D) Leskea polycarpa (D) Dianthus microlepis {O)

ASPLENIETEA TRICHOMANIS

Carex kitaibeliana Helianthemum canum Tortella tortuosa Euphrasia salisburgensis Achillea * aizoon Thymus cherlerioides Campanula *�pirinica Encalypta streptocarpa Asplenium ruta-muraria Leontopodiurn * nivale Genista albida Jurinea glycacantha Sedum dasyphyllum Kernera saxatilis Sedum oc.hroleucum Seseli rigidum Asplenium trichomanes Brassica jordanoffi Iberis pruitii Silene saxifraga

FESTUCO-SESLERIETEA

Saxifraga ferdinandi-coburgi Festuca * pirinica Sesleria coerulans Anthyllis vitellina

22

....... 2. 5583757

... 32 ..

33 ... 3.

2 .. 22 ..

. 22.2 ..

. 22 ....

55335.255 6634464

323355325 33.3433

3.232.532 322.534

3233332.3 .33 .. 42

333254333 .2.3.2.

322 .. 23.2 232 .. 23

... 2 ... 33 222 ... 2

...... 343 22 .. 4 ..

2.22.222.

.. 2 .... 2. 22.2 ...

..... 3 .. 5 .. 2.2 ..

.... 2 .. 22 .2 .... .

...... 222 ..... 1.

...... 22.

.. 2 ... 3 ..

.. 3 .. 2 ...

....... 2.

....... 3.

........ 2

........ 2

35333 .. 23 344.454

43332433. 3324 ...

3 .. 6.4634 .42.323

.22 .. 2.22 222.22.

Cl C2

% %

100' 29

100 14 89 29

78 14 78 14 67 O

67 O 67 O 89 14 67 O 67 14 56 O 56 O 44 O 44 O

33 O 33 O

11 100

O 29

O 43

O 43

O 43

O 29

89 100

100 86

78 86

89 57

100 43

67 71

33 57

33 43

67 O 22 43

22 29

33 14

33 14 22 O 22 O

22 O

11 O 11 O 11 O 11 O

78 86

89 57

67 71

56 71

Androsace * arachnoidea Aster * dolomiticus Hieracium bifidum Oxytropis urumovii Poa alpina Acinos alpinus Calamagrostis varia Carex sempervirens Centaurea achtarovii Koeleria eriostachya Linum extraaxillare Saxifraga paniculata Scutellaria alpina Sedum annuum Silene * graefferi Silene * pirinica

THLASPIETEA ROTUNDIFOLII

Saxifraga luteoviridis Thlaspi bellidifolium Viola grisebachiana Bromus * lacmonicus Thalictrum minus Arabis ferdinandi-coburgi Senecio rupestris Saxifraga exarata Galium stojanovii

SALICETEA HERBACEAE

Sedum atratum Dryas octopetala

ERICO-PINETEA

Daphne oleoides Pinus leucodermis juv.

OTHER PHANEROGAMS

Campanula velebitica Sonchus sp. Euphorbia cyparissias Campanula rapunculoides Alyssoides graeca Pedicularis sp. Scabiosa cf. webbiana Sempervivum arachnoideum Thymus sp. Carduus scardicus Epipactis helleborine Juniperus * nana

OTHER CRYPTOGAMS

Ditric�um flexicaule Hypnum cupressiforme Psora decipiens Squamarina gypsacea Toninia coeruleonigricans Collema sp. Peltigera rufescens Solorina bispora Cirriphyllum tenuinerve Tortula muralis Myurella jullacea Timmia bavarica Cladonia pocillum Schistidium apocarpum Ceratodon purpureus Grimmia tergestina Fissidens cristatus Pseudoleskea nervosa Dermatocarpon miniatum Endocarpon miniatum Placynthium nigrum

....... 25 22 .... 3

... 2 ... 2 . . 2 ... 21

... 22.22 . ...... 2 332... . . . . ..... 2 ..... 3 .. 2 .. 2. 2 .. ...... 2. . . ... 2. 2 2 ....... . ..•..••• 5

...... 2 .•

....... 2.

...... 2 ..

..... 3 ... •....... 2

. 2 ......•

..... 2.

.•.• 2 ••

,2 .. 23332. 123 .. 2. 333 ... 3 .. ...... 22. . ... 2. 2

... 332 ..... 2. 22

2 ...... 2. ... 2 .... .

2.

..... 2. •. 2 ....

..... 1. •.... 5.

... 2 ... 3. . .... 3.

. . 2 .. 1. ..

. 2. 2. 3 .. 2. 4 ... . 2 ..... 2. 2. . . . 2.

... 2. ... 2.

.. 2. ... 2. . 2.

.1 .. 1..

. . . . . 1.

.. 2. 2 .. 4. 22 ..

33 ...... 4 . 4 ... 3

. 3 ...... 2 . 2 ..••

.. 22

..... 2. 2.

.. 3 .. . 2 ..

.... 3 .... ...... 3.

. 2 ..•...•. 2 ........ 2 . 2 ...... .

.. 2

. 2.

2.23222 2. 22 ...

.... 4. 2

... 23. 3

..• 3 •.• 2 ...•..

.. 2 ....

... 2 ..•

.•. 2 ••.

.. 2 •.•• .. 2.

22 43

22 43

22 14 33 14

22 29

11 29

11 O

11 O

O 14

o o

11 O

11 O

11 O

11 O

O 14

11 O

67 57 44 O

22 29

33 O 33 O 22 O 11 O

O 14

O 14

11 14

O 14

22 14

22 O

33 22 22 22 11

11

11

11 11

11 11

11

33 22 33

22 22

11

22 22

o o o o o

11

11

11

11

11

11

11

11

o o o o o o o o o o o o

86 43 29

43 14

14 o o

14

14

14 14

14

o o o o o o o o

Notes: C - constancy; Ch - character species; D - differential species; * - subspecies.

23

Tab. 2 -Asplenietea trichomanis of the Balkan Peninsula (a short version). The species importance is given in constancy classes according to Braun-Blanquet (1964).

No. of source table No. of relevés per table

2 10

3 14

ASSOCIATIONS OF THE RAMONDION NATHALIAE ICh & Dl:

Viola kosaninii V Saxifraga scardica V II

Potentilla speciosa V

Minuartia graminifolia III I

Asyneurr.a limonifolium II

Saxifraga JT1arginata IV

Arenaria eretica III

Oraphne alpina

Jurinea consaguinea III Inula . aschersoniana II

Sax i fn1ga juniperifolia

Hypochoeris • pelivanovicii Asperula pirinica Centarea mannagettae Globularia meridionalis Allium . minus

Saxifraga . brevifolia

RAMONDION NATHALIAE

Carex kitaibeliana fDl I II III

Silene saxifraga agg. (Chl II III III I I Hieracium pannosum IChl II II III

Ramonda nathaliae IChl V III V V

Saxifraga grisebachii IChl III III IV

Campanula formanekiana (Ch) V V

Saxifraga porophyla ICh) I IV

Alyssum corymbosum (Ch) I I

Frangula rupestris (D) I I III III .

Draba elongata IDJ II II I

Campanula versi color IDI II III

Minuartia verna (Ch) IV

Achillea ageratifolia agg. (Ch) II

Potentilla . stojanovii (Ch)

Leontopodium * nivale ICh)

Campanula . pirinica ICh)

ASSOCIATIONS OF THE EDRAIANTHO-ERYSIMION ICh & D)

Ramonda serbica Alyssum . orientale

Viola grisebachiana Thymus serbicus

EDRAIANTHO-ERYSIMION COMATAE

Carum rigidulum (incl. graecum) Sedum ochroleucum Campanula rotundifolia Dianthus petraeus Aster alpinus Cincl. dolomiticus)

Sesleria rigida IDI Erysimum comatum IChJ"

Edraianthus graminifolius IChl

Cerastium banaticum (Ch)

Edraianthus serbicus ICh)

24

III

V

V

IV IV

IV

V IV V II

I V

V I II V

III V I

IV III I II II II

II II

III V

I I

IV II

III

III II

9 20

V

I

III

V

II

IV

IV

III

10 10

V

V

IV

IV

IV

V

V

V

11 10

V

IV

II

IV

IV

V

I

v.

IV

II

III

12 5

V

IV

IV

IV

II

IV

IV

II

I

II

13 2

2+-1 2+-1 2+ 2+-1

2+-1 2+

2+

2+-1

POTENTILLETALIA CAULESCENTIS

Micromeria cristata IV II V III IV II

Seseli rigidulum IV I IV II IV

Asplenium viride III III I 2+-1

Potentilla . apennina V IV 22-3


Asplenium ruta-muraria III IV III V III II IV IV 2+

Asplenium trichomanes I I II I IV IV II

Sedum dasyphyllum IV I III III II

Silene pusilla I II III I 2+-1

Asplenium fissum I I II

Draba aizoides I III

Cystopteris fragilis IV I

Localities of tables (l'ab. 2):

1. Micromerio-Violetum kosaninii; Horvat et al. (197 4), Tab. 140, column 1; Macedonia

2. Saxifrago-Potentilletum speciosae; Horvat et al. (1974), Tab. 140, column 2; Galicìca, Bistra,

Korab (l\tlacedonia)

3. Saxifrago karadzicensis-Potentilletum apenninae; Horvat et al. (1974), Tab. 140, column 3;

Jakupica (l\tlacedonia)

4. Campanuletum formanekianae; Horvat et al. (1974), Tab. 140, column 4; Nidza (l\tlacedonia)

5. Campanula formanekianae-Ramondietum nathaliae; Quézel (1967), Tab. 6; Vermion (Greece)

6. Leontopodio-Potentilletum stojanovii; Simon (1958), Tab. I; Pirin (Bulgaria)

7. Leontopodio-Potentilletum stojanovii; Tab. 1 in this study, relevés 10-16; Pi.rin

8. Hieracio pannosi-Caricetum kitaibelianae; Tab. 1 in this study, relevés 1-9; Pirin

9. Saxifragetum brevifoliae; Horvat et al. (1974), Tab. 140, column 6; eastern Serbia

10. Erysimo-Ramondietum nathaliae; Jovanovié-Dunjié (1953), Tab. 1; Suva Planina (Serbia)

11. Ceteracho-Ramondietum serbicae; Jovanovié-Dunjié (1953), Tab. 2; Rtanj (Serbia)

12. Saxifrago aizoi-Violetum grisebachianae; Jovanovié-Dunjié (1956), Tab. 12; Rtanj (Serbia)

13. Potentilla apennina-Saxifraga aizoon-Ges .; Jovanovié-Dunjié (1955); p. 88; Suva Planina (Serbia)

Floristically and synecologically analogous community was reported from the BucegiMts. by Domin (1983). This community grows on gentle slopes covered with water-soaked soils on limestone. It is frequented by dominant Silene pusilla and Doronicum carpaticum; further Saxifraga paniculata, S. adscendens, S. androsacea, Viola biflora and Mysotis alpestris are also found. The community is also rich in mosses.

Syntaxonomy of the Asplenietea trichomanis communities

The plant communities in rock fissures and crevices of the Balkans ever enjoyed much interest of vegetation systematists. Major motivations for this interest are the high number of endemie and relict species found in the communities as well as diversity of types encoutered over the varied high-mountain landscape of the Balkan Peninsula. The syntaxonomy of the Asplenietea trichomanis in the Balkan Peninsula is based mainly on works ofHorvat (1931, 1934, 1936, 1937, 1960) who described many types and several high-ranked syntaxa from Yugoslavia (Macedonia, Serbia, Croatia). Quézel (1964, 196 7) described rock-fissure communities from

25

Tab. 3 - Silene pusilla-Saxifraga oppositifolia community.

No of relevé

Exposition

Slope 0

Sampled area rn2

Cover of herb layer %

Cover of moss layer %


Silene pusilla Campanula·• pirinica Asplenium fissum Encalypta streptocarpa Euphrasia salisburgensis Thymus cherlerioides


Galium stojanovii Saxifraga oppositifolia Veronica * kellereri Viola grisebachiana Alyssum * pirinicum Armeria * alpina Myosotis alpestris Thlaspi bellidifolium Papaver * degenii Doronicum columnae Saxifraga * discolor Saxifraga * pirinica Arabis caucasica Saxifraga luteoviridis

FESTUCO-SESLERIETEA

Cerastium * lanatum Poa alpina Pedicularis verticillata Saxifraga ferdinandi-coburgi

SALICETEA HERBAC�AE

Poa pirinica Draba scardica Omalotheca supina Sedum atraturn

OTHER PHANEROGAMS

Hieracium sp. Taraxacum sp.

CRYPTOGAMS

Leskea polycarpa Hymenostylium recurvirostre

26

12

ww

ss

ww

89

00

94

12

55

<3 55

56 55 23

22

11

.2

36

23

23

23

22

22 22

22

21

6. 2. 3. • 2

.1

22 22

.1

.1

22

• 2

• 3

• 2

.2

.2

37

35

Tortella tortuosa Bryurn caespiticiurn Collerna sp. Toninia sp. Tortula sinensis

Bryurn elegans Protoblastenia terricola

Psora decipiens Hypnurn cupressiforrne Leptogiurn sinuaturn Plagiochila porelloides

33 22 22 22 22 3. 2. 2.

. 2 • 2

. 2

Greece. Many papers deal with the Asplenietea trichomanis in Rumania (eg. Borhidi 1958, Schneider-Binder 1968, 1969, 1972, 1975, 1980, Schneider-Binder & Voik 1977, Tau ber 1985). In Bulgaria Pawlowski et al. (1937) and Simon (1958) brought descriptions of same new associations of the Asplenietea trichomanis.

The communities typical of lime-rich substrata �imestone, marble) from Macedonia and Bulgaria were classified within the Ramondion nathaliae (cf. Horvat 1936, Jovanovié-Dunjié 1953, 1955, 1956, Simon 1958, Bleèié & Tatié 1960).

Ramondion nathaliae suggested Horvat (1935, 1936) for south-Macedonian mountain ranges. The descriptions are, however, rather unclear. No diagnostic species ate given and the associations supposed to be included within the Ramondion nathaliae are also not listed. Horvat (1936) listed only a few species, such as Ramonda nathaliae, Saxifraga scardica, S. marginata, S. grisebachii, Potentilla apennina, Campanula formanekiana, Micromeria cristata and some others, as dominants. The same name was used by Jovanovié-Dunjié (1953, 1955, 1956) and Bleèié &Tatié (1960) for a group of east-Serbian communities.Neither in these papers the validisation of the alliance was performed. Ramondion nathaliae was validated by Simon (1958) by listing character species of the alliance (see Tab.I in Simon 1958). He assigned the Leontopodio - Potentilletum stojanouii into the Ramondion nathaliae sensu Horvat (1935), thus interpreted the alliance as that comprising the fissure plant communities occurring in southern Macedonia (Yugoslavia);Bulgaria and most probably also some mountains of northern Greece and Albania. Quézel (1967) listedMinuartia uema, Saxifraga grisebachii (also near Bansko; Kuzmanov 1970), Micromeria cristata and Achillea agerati/olia subsp. aizoon as members of the diagnostic species group for the Ramondion nathaliae. Horvat et al. (197 4) added also Hieracium pannosum, Campanula uersicolor, Alyssum corymbosum and Cerastium banaticum to the diagnostic species of the alliance.

According to our opinion the use of the Ramondion nathaliae for the eastSerbian communities is incorrect because of the floristic differences between the Macedonian and Serbian plant communities oftheAsplenietea trichomanis (Tab. 2). Therefore we describe a new alliance for the east-Serbian communities, the Edriantho graminifolii - Erysimion comatae all. nova (nomenclatura! type: Ceteracho - Ramondietum serbicae J ovanovié-Dunjié 195 2). The diagnosti e taxa of the alliancecomprise Erysimum comatum (char.), Edraianthus graminifolius (char.), E. serbicus(char .) , Cerastium banaticum (reg. char .) , Sesleria rigida (dif .), Sedum ochroleucum(char.) Ramonda serbica (char.), Alyssum saxatile subsp. orientale (char.), Violagrisebachiana (reg. char.) and Thymus serbicus (char.).

27

It should be noted, however, that the Bulgarian communities (Leontopodio Potentilletum stojanovii and Hieracio pannosi-Caricetum) are differing from the Yugoslavian Macedonian group by the endemie Bulgarian (and Pirin) species, and at the same time show some floristic relations to the Edraiantho-Erysimion comatae by common occurrence of species such asAster alpinus, Carum rigidulum agg., Campanula rotundifolia, Dianthus petraeus, Sedum ochroleucum and the like.

According to the syntaxonomic revision summarized in Tab. 2 we suggest the following classification of Potentilletalia caulescentis in southeastern Balkan Peninsula:

Asplenietea trichomanis (Br. - Bl. in Meier et Br. - Bl. 1934) Oberd. 1977 Potentilletalia caulescentis Br. - Bl. 1926 Ramondion nathaliae Horvat ex Simon 1958

1. Micromerio-Violetum kosaninii Horvat ex Horvat et al. 19742. Saxifrago-Potentilletum speciosae Horvat et al. 197 4 (syn. Potentilla speciosa

Minuartia graminifolia-Ass. Horvat 1937)3. Saxifrago karadzicensis-Potentilletum apenninae Horvat ex Horvat et al.

19744. Campanulo-Inuletum candidae Horvat 19495. Campanuletum formanekianae Horvat 19386. Campanula formanekianae-Ramondietum nathaliae Quézel 19677. Achilleo-Aubrietetum gracilis Horvat 19368. Leontopodio-Potentilletum stojanovii Simon 19589. Hieracio pannosi-Caricetum kitaibelianae Mucina et al. 1990

Edraiantho graminifolii-Erysimion comatae Mucina et al. 1990

10. Erysimo-Ramondietum nathaliae Jovanovié-Dunjié 195211. Saxifragetum brevifoliae Bleè'i.é et Tatié 196012. Ceteracho-Ramondietum serbicae Jovanovié-Dunjié 195213. Saxifrago aizoi-Violetum grisebachianae J ovanovié-Dunjié ex Mucina et al.

hoc loco(syn. As. Saxifraga Aizoon-Viola Grisebachiana prov.; Jovanovié-Dunjié1956)nomenclatura! type: Tab. 12, relevé 4 in Jovanovié-Dunjié (1956, p. 34),lectotypus

14. Potentilla appennina-Saxifraga aizoon community (Jovanovié-Dunjié 1955)

3.2. Plant Communities on Marble Screes

High altidudes, both in the subalpine �ow-situated sites in glacial valleys) and alpine (slopes of high-elevated glacial circles) belts in the marble part of the Northern Pirin Planina Mts. are rich in scree habitats resembling those in the Alps (Zollitsch 1966). Three associations, including the Papaveri degenii-Armerietum

28

alpinae, Veronico kellereri-Silenetum prostratae (belonging to a new alliance, the Veronico-Papaverion degenii), and the Bromo lacmonices-Geranietum macrorrhizi (the Silenion marginatae) are described in the sequel.

Papaveri degenii-Armerietum alpina e ass. nova Nomenclatura! type: Tab. 4, relevé 12,hoc loco

The Papaveri degenii-Armerietum alpinae is a pioneer community of low-grown herbs, grasses and mosses growing on crystalline limestone screes in the alpine belt of the Pirin Planina Mts.

The stands populate foothill screes and slopes of moraines at altitudes between 2200 to 2700 m. The slope of the screes ranks between 25-45° and only exceptionally are more gentle (5-10°); aspect of the slopes is prevailingly north and northeastern, to a lesser extent western or southwestern. The substratum is formed by rocks having 10 to 20 cm in diameter and intermingled with gravel (3 to 5 cm in 0). Fine -grained soil particles occur in varying quantities. Most of the habitats are well-supplied by water which comes either directly from rainfall or, mainly, from thawing snow and firn patches. Large water streams emerging in spring usually give rise to erosion dells in the scree cones, and aid, together with large boulders, differentiation of the scree into variety of microhabitats. This reflects also in high species richness of the community.

The stands, sometimes more than 50 m2 large, and with total cover ranging from 2 5 to 7 5 % , are dominated by chasmophytes. The stands are bistra tal; the lower herb sublayer (1 to 5 cm) is formed by dwarf Arenaria biflora, Silene pusilla, Cerastium alpinum subsp. lanatum, Galium stojanovii, Saxifraga appositi/olia and rosettes of several Saxifraga species. The higher herb sublayer (5 to 15, max. 25 cm high) is formed by the dominants ofthe community, such asArmeria pocutica subsp .alpina, Doronicum columnae, Papaver degenii, Myosotis alpestris, and Veronica saturejoides subsp.kellereri. The moss layer (1 to 4 cm) is also well-developed, mainly in wetter microhabitats, where it may attain 30 to 60%. The most frequent cryptogams include Ditrichum flexicaule, Hymenostylium recurvirostre, Leskea polycarpa, Tortella tortuosa, Preissia quadrata and Tortula sinensis.

Syndynamically the Papaveri-Armerietum is seen as arrested (blocked) successional stage (sensu Moravec 1969) of a pioneer character. In dried and more insolated habitats, after the screes has been partly consolidated by fine-grained soil, the stands be come impoverished of typical sere e species (Veronica saturejoides subsp. kellereri, Doronicum columnae, Saxifraga adscendens subsp. discolor) while tussock-forming grasses and sedges, such as Sesleria coerulans, Festuca riloensis (only rarely), Carex rupestris, Carex laevis (frequently) take over. The further succession proceeds in direction to the Anthyllo-Seslerion klasterskyi Simon 1958 (Festuco-Seslerietea Barbero et Bonin 1969), most probably towards the Carici rupestris-Seslerietum klasterskyi Simon 1958 described from the Vikhren Mt. The latter community forms a transition between the Veronico-Papaverion degenii and Anthyllo-Seslerion klasterskyi. In wetter, shady and gentle-sloped habitats the

29

Tab. 4 -The communities of the Veronico-Papaverion degenii.

No. of relevé

Exposition

Slope 0

Sampled area mz



PAPAVERI-ARMERIETUM ALPINAE

Saxifraga • discolor (Ch) Papaver • degenii {Ch) Viola grisebachiana (Ch) Saxifraga androsacea (Ch) Ditrichum flexicaule ID) Leskea polycarpa (D) Sedum atratum (O) Pedicularis verticillata (D) Saxifraga exarata (Dl Draba scardica {D) Peltigera rufescens (Dl Solorina bispora {Dl

D-TYPICUM

Aren«ria biflol'a Omalotheca supina Hymenostyliurn recurvirostre Preissia quadrata Mnium stellare Salix reticulata Primula minima Carex parviflora Drepanocladus aduncus Plantago atrata

D-FESTUCETOSUM RILOENSIS

Festuca riloensis Alyssum • pirinicum (O) Pedicularis orthantha Dianthus microlepis

VERONICO-SILENETUM PROSTRATAE

Silene • prostrata (Chl Senecio rupestris ID) Festuca valida (Dl Linum capitatum (Dl

VERONICO-PAPAVERION DEGENII

Poa pirinica Veronica • kellereri Galium stojanovii Erigeron vichrenensis Arabis ferdinandi-coburgi


Doronicum columnae Armeria • alpina Myosotis alpestris

11111111112222 222222

12345678901234567890123 456789

NN N NN EE N NN W EE EE

NE NNNN NN N NNNSSNN sss s

ENNEEWENEENENWNEEEWWEEN EEEESE

3143313133433 342334441 444444 00500005050505555505005 000505

1 1 253232 312233134 486553 60885331058586001450650 803006

46255253533325446433531 243432 00005505055050000000505 000050

365<6312< 1<5 <l< « « < <

000100505-55555505--15- 15-1-1

2.32 .. 255333333233232.3

.2352 .. 255225 ... 2.55355 2 .....

22.22 ... 2222.2.2523522.

775775552 ........ 2 .... .

23 .. 3552.2322633233 ... .

542.62.3 .. 54.5 .. 3 ... 2 ..

....... 2222.2222 .... 21.

... 122 ... 22.2 .. 21.

.... 33 ...... 2 ..... 35.2.

... 2 ......... 2·.1 .. 2 ... 2

...... 2. .23.2 ..... 1.

.. 2.22 ...... 2 ....... 2 ..

33232.2222.2255232 ... . .3.222

22 ... 2222.23.52535 .... .

67 .. 7343 .. 53 ... 22 .... . . 2 ....

22 .... 3.2.22 .. 32.2 .... .

24 .. 22 .. 2 .. 2.32 ....... .

23 ..... 5 ..... 22 .. 2.

.. 2.53 ..... 3 .. 2 ...... .

. 2. 2. . ...... 5 ........ .

. . 8 .. 7.

. 2. . . .... 3 .......••

... 3 ... 5253.

.. 3553 . . 2.2.5

... 2. 2 .. 2. 2.

.... 2 .. 2 .. 4. . 2 ..

.....•........ 2. 276776

............... 223.22

.....•.....••...... 3 ... 5426.3

· · · · · · · · · · · · · · · · · · · · · · · .2.522

522353332553255355 .. 33. . 4. 34. 22 .... 2.33523325532222. 363655

3 ... 2 .... 2322335355. 33. 2.2322

......... 2 .. 2222 ...

........ 2.2 ....... . .. 4. 3

355.35775555577352.353. 55523.

32332.2253.555327755755 22. 2 ..

23232525755553555523.25 5 ... 4.

30

Cl

%

83

70

70

43

70

48 43

35

26

22

22

22

70

57

39

39

35

26

22

13

9

9

22

17

13

13

4 o

4 o

87

74 61

22

9

87

91

96

C2

%

o

17

o

o

o

o

o

o

o

o

o

o

67

o

17

o

o

o

o

o

o

o

o

50

17

17

100

83

83

67

50

100

83

o

33

83

50

50

Saxifraga oppositifolia Arabis caucasica Saxifraga luteoviridis Thlaspi bellidifolium Tortula sinensis Myosotis suaveolens Silene acaulis Geum reptans Veronica alpina Scrophularia * laciniata

FESTUCO-SESLERIETEA

Poa alpina Sesleria coerulans Cerastium * lanatum Helianthemum * tomentosum Botrychium lunaria Saxifraga ferdinandi-coburgi Ac-inos alpinus Silene * graefferi


Silene pusilla Tortella tortuosa Euphrasia salisburgensis Cystopteris regia Thymus cherlerioides Carex kitaibeliana Potentilla * stojanovii Campanula * pirinica Helianthemum canum Asperula suberosa Carum rigidulum agg.

SALICETEA HERBACEAE

Ligusticum mutellina Bartsia alpina Gentiana verna Sagina saginoides Saxifraga heucherifolia Potentilla crantzii Anthelia juratzkana Artemisia eriantha Polygonum viviparum Primula halleri Soldanella pusilla

OTHER PHANEROGAMS

Taraxacum nigricans agg. Hieracium Bg24 Juniperus * nana . Knautia sp. Cerastium sp.

OTHER CRYPTOGAMS

Bryum caespiticium Brachythecium velutinum Bryum elegans Collema sp. Encalypta streptocarpa Cladonia sp. Toninia candida Anomodon viticulosus Bryum sp. Catopyrenium cinereum Cirriphyllum cirrosum Cladonia pocillum Collema tenax Hypnum cupressiforme Leptogium lichenoides Plerospora hookeri Pseudoleskea nervosa Tortula intermedia Timmia austriaca Jungermania leiantha

235232232225523.23535.5 253222.2235222.2 ...... . 2 ...... 1.1222.2222.2 .. . ........ 232 .... 55.33 .. 2 2 ... 62.5 .... 2 ... 2 ..... . .............. 2 ... 2.3 .. ........... 3 .. 3 ... 5 ... 2 .•.... 5 ...... 52 ....... . ••............ 5 •• 2 .••••

2 .... . 3 .... . ... 2 .. ... 232 22 .... . . 42. 3

••••• 2

3 .. 2 .. 2.3.2.3 .. 2.235355 5 ..... 54 ........ 23.2.5532.2.2 2 .. 422 .. 22 .... 22 .. 32 .... 55.23 ... 2 .. 3. . . . . . . . . . . . . . . . . . . . . . . .. 5. 2 ............... 25 .... 2. ........... 2 ... 22 ..... . ...••.................. .•. 3 •. ..••................. 5. . .. 2 ..

23.2 .. 22.233225323 ... 2. 2 ....... 2222.2.32 ..... .

....... 1. ...... 2. 22 ........ 2 .... 3 ...... . .............. 2.· ..... 2. ............... 3.3 .... . .............. 1. ...... .

. 2 ......... 2 ... .

..... 2 .•.. ......•...... 3 ...••.••• . 3.

.....•.... 2 ... ..... 2 .•.••.•

.............. 2 ... . .2.... . ........... . ............... 1 ...... . ....... 5 .............. .

. 2. 2. 2

. .. 2. 2

... 2. 3 ••• 2 •• ... 2 .. ... 2 ..

.... 23

55232.235332.3255522232 55223. . . . . . . . . . . . . . . . . . . . . . . . . .. 22. ....................... ... 1 ..

.••.•.•.......•..••••• ••• 2 •• . .••...... 2 .......•....

.. 2. 2. 5 ............. 2.

.... 22 .. 22 ......... 2 .. 2 ..... 3 ......... 2 ..... . 2 ..... 2 ...... 2 ........ . 2 .... 3 ......... 2 ......• . 2 ........... 3 ........•

. 22 ................... .

..•.•. 6 ..............•. ............ 3 ......•••

............. 2 .......••

.... 2 ................. .

....... 2 .............. . ...........•...... 2.

•••....• 3 .•...•..•..... 2 ..................... . ....... 2 .......... .

..............• 5 .•..•••

.. ·.· ...... 2 ........... .

.6 .................... .

.2 .................... .

Notes: Cl - constancy of Papaveri-Armerietum; C2 - contsancy of Veronico-Silenetum prostratae; for other abbreviations see Tab. 1.

31

91 61 48

35 26 13 17 13

9

o

57 48

43

4

13 13

o 4

65 35

9

17 9 9 4

o

o

o

o

o

9 4

4

4

4

4

4

4

4

4

91 o o o 4

17 22 13 13 13

9

13 4 4

4

4

4

4

4 4 4

4

4

4 4

17 17 17 50 33 50

o o o

17

17 67 17 33

o o

17 17

o

50 33

o

o

o

o

33 17 17 17

33 o

o

o

o

o

o

o

o

o

o

83 33 17 17

o

o o o o o o o o o o o o o o o o o o o o

development might proceed towards the Salicetea herbaceae, as also shown by the occurrence of many snow-patch species in the subassociation typicum.

The Papaveri -Armerietum is a rather common feature of scree slopes of all of the glacial circles in the marble part of the Northern Pirin Planina Mts. It is supposed to be locateci also at the Sinanitsa Mt.

Papaver degenii, Saxifraga adscendens subsp. discolor, S. androsacea and Viola grisebachiana are the character taxa of the association, while Ditrichum flexicaule, Leskea polycarpa, Sedum atratum, Solorina bispora, Pedicularis verticillata, Saxifraga exarata subsp.pirinica, Draba scardica and Peltigera rufescens differentiate the Papaveri-Armerietum well from the Veronico-Silenetum prostratae (see below).

In relation to soil moisture, the degree of scree stabilization and granulometric composition of the soil, the Papaveri-Armerietum is differentiated into a series of synecologically and floristically well-discernible units including the subassociation typicum (with typical variant and variant with Saxifraga androsacea) and the festucetosum riloensis.

Papaveri degenii-Armerietum alpinae typicum subass. nova Nomenclatural type: identical with that of the association.

The soil supporting this community is skeletal, rich in boulders and gravel, and with a varying amount of fine-grained materiai. The slope of the sites is frequently 30 to 45°. among aspects north is the prevailing one, while east and west occur less frequently. The community occurs in extensive stands (up to 100 m2);they are 10 to 25 cm high and poor in mosses. The differential species of the subassociation are

Arenaria biflora, Omalotheca supina, Salix reticulata, Primula minima, Carex parviflora, Plantago atrata, Preissia quadrata, Hymenostylium recurvirostre, Mnium stellare and Drepanocladus aduncus.

There are two synecologically well-interpretable variants within the typical subassociation such as the variant with Saxifraga androsacea and typical variant. The stands of the former one populate the wettest habitats soaked with percolating water coming from thawing snow patches in upper positions of the scree cones found under steep rocky walls in mouthes of rocky dells and gorges. The finegrained to gravelly, largely stabilized sere es (1 O to 30° steep) face north or northeast. The stands are carpet-like, low-grown (5 to 10 cm) and small-sized. At sites with gentle slope these habitats resemble snow patches. The community is characterized by dominance of Saxifraga androsacea, which seems to occur more frequently on silicate bedrock in other mountain ranges of Europe (eg. Domin 1930). The high cover values of several moss species such as Drepanocladus aduncus, Hymenostylium recurvirostre, Leskea polycarpa and Tortula sinensis are also of diagnostic value for the variant.

Along a complex ecocline (from wet habitats with fine-grained soil towards sunny and dried habitats with coarse-grained screes) the typical variant occupies an intermediate position between the Saxifraga androsacea variant and the subassociation festucetosum riloensis.

32

Papaveri degenii-Armerietum alpinae festucetosum riloensis subass. nova Nomenclatura! type: Tab. 4, relevé 21 hoc loco

The subassociation is limited to well-insolated habitats enjoying all-day-long sunshine. These are found on convex scree con es and ridges of moraines. The screes are gravelly, with a share of small rocks; the amount of fine-grained material and boulder is very varying. The slopes housing stands of the community usually face south-west to east-north or east aspects. The community occurs frequently at altitudes between 2600 to 2700 m. The are stands rather extensive and only poorly covered (35% on the average). Mosses are rare. The differential taxa of the subassociation include Alyssum cuneifolium subsp. pirinicum, Festuca riloensis, Dianthus microlepis and Pedicularis orthantha. The latter species occurs more frequently on granitic bedrock especially in central and southem parts of the Pirin Planina Mts. or in other Bulgarian mountain ranges, eg. Rila Planina Mts. where it is considered a character species of the Carici curuulae-Festucetum riloensis Horvat et al. 1937 (Seslerion comosae Horvat 1935, Caricetea curuulae Br. - Bl. 1948).

In comparison to other Veronico-Papauerion communities, the festucetosum riloensis is characterized by higher frequency of occurrence of Poa alpina and Cerastium alpinum subsp. lanatum.

Veronico kellereri-Silenetum prostrata e ass. nova Nomenclatura! type: Tab. 4, relevé 27,hoc loco

The Veronico - Silenetum prostratae is a pioneer community of herbs populating bouldery marble screes in the alpine belt of the Pirin Planina Mts.

The community was recorded from the Golemiya Kazan Circle on moving screes found on south and southeast-facing steep slopes (40 to 45°). The screes were built of marble skeleton (5 to 30 cm in 0) and admixed boulders as large as 1 m in diameter. A small amount of fine-grained eroded rocky material flushed on the bottoms of deep cracks among the boulders is also found. The screes are insolated during the most of the day, thus the sites are relatively warm and dry.

The stands of the community are 50 to 100 m2 large, with rather loose canopy (cover 20 to 40% of a plot) and are strikingly stra tifi ed. The upper-most sublayer (up to 30 cm) is formed by the dominant Silene uulgaris subsp. prostrata and accompanied by Doronicum columnae. The middle herb sublayer (10 to 20 cm) is the species-richest stratum of the stands. Veronica saturejoides subsp. kellereri,

Senecio rupestris, Armeria pocutica subsp. alpina, Linum capitatum, Myosotis

alpestris, M. suaueolens and Arabis ferdinandi-coburgi are found in it. The lower herb sublayer (up to 5 cm) is frequented by dwarf-grown species such as Galium stojanouii, Arabis biflora andEuphrasia salisburgensis. Due to extreme microclimatic conditions and the overall dryness of its sites the community is poor in mosses.

Silene uulgaris subsp .prostrata occurs in highmountains on calcareous screes in many European countries, where it is considered a typical scree element (Hadaè' et al. 1969, Duvigneaud et al. 1970, Richard 1971, Valachoviè' 1989).

33

The dominant taxon, together with Festuca valida, Senecio rupestris and Linum capitatum form the character taxon combination of the Veronico-Silenetum pro

. stratae. A synecologically analogous community of lower altitudes is the Bromo lacmonices -Geranietum macrorrhizi (see below). The following relevé is a transition between these two units:

Kamenititsa Tsirkus, alti tu de 2200 m, aspect E, slope 3 0°, sampled area 30 m 2, cover of herb layer 40%; August 13, 1984.

Bromus cappadocicus subsp. lacmonices 1, Scrophularia heterophy/la subsp. laciniata 2a, Senecio rupestris 1, Silene vulgaris subsp .prostrata 1, Thalictrum minus su bsp. olympicum 3 ,Arabis caucasica 1,

Doronicum columnae +, Festuca valida +, Mysotis alpestris +, Veronica saturejoides subsp. kellereri +,

Sesleria coerulans 2a,Poa alpina 2 m,Acinos a/pinus 1,Cirsium appendiculatum 1,Hieracium grandif/,orum 1, Onobrychis montana su bsp. scardica + ,Dianthus scardicus + ,Phleum montanum +, Euphorbia cyparissias +,E. amygdaloides +, Galium anisophyllum agg. + ,Asplenium fissum + ,Kemera saxatilis +,

Hieracium pannosum r. Scorzonera rosea +, Daphne oleoides +, Taraxacum sp. +.

Bromo lacmonices -Geranietum macrorrhizi ass. nova Nomenclatura! type: Tab. 5, relevé 3 hoc loco

The community populates moving or slightly stabilized screes on east- and south-facing slopes (15 to 40°) that occur mainly in the belt of Pinus leucodermis (at altitudes between 1800 to 2000 m). The skeleton building the screes is the marble stones (20 to 30 cm in diameter); also large boulders (up to 1 m in diameter) are present. At lower altitudes, also an admixture of granite stones was observed. Gravel and fine-grained material is scarce and it is concentrated on bottoms of the fissures among the sere e rocks.

The stands of the community are from 20 to 100 m2 in extent. U sually they are covering the sere e in broad belts following the main axis of the scree. The total coverage is dependent on the cover of Geranium macrorrhizum, a chasmophyte broadly tolerating the range of soil reaction. Other species occur sparsely. Some epiterrestric mosses occur among the boulders. More frequently occur epilithic lichens and mosses.

The group of character taxa consists of Geranium macrorrhizum, Bromus cappadocicus subsp. lacmonices and Moehringia pendula. The classification of the Bromo-Geranietum macrorrhizi is problematic because of the broad habitat requirements of the dominant species (Bo�caiu 1971, Sanda et al. 1977). Geranium macrorrhizum is distributed (in Europe) in the Apennines, Southern Alps, Eastern and Southern Carpathians and other Balkan mountains. Most often it is limited to calcareous screes. In southern Greece (the Parnassos Mts .) it is a dominant species intheSenecioni thapsiformi -Geranietum macrorrhizi (Quézel 1964),a vicariant unit to the Bromo-Geranietum macrorrhizi, occurring in analogous habitats and in the same altitudinal belt. Quézel O.e.) classified the community within the Silenion caesiae Quézel 1967 (Drypetea spinosae Quézel 1964). However, both associations share only one species - Geranium macrorrhizum. In Southern Alps, Geranium macrorrhizum occurs in the Moehringio-Gymnocarpietum robertiani (Petasition paradoxi Zollitsch 1966,Drabetalia hoppeanae Zollitsch 1966, Thlaspietea rotundi-

34

Tab. 5 - Bromo-Geranietum macrorrhizi (Silenion marginatae).

No. of relevé

Exposition

Slope o

Sampled area m2


BROMO-GERANIETUM MACRORRHIZI

Geraniurn macrorrhizum (Ch) Bromus * lacmonices (Ch) Moehringia pendula (Ch)


Thalictrum * olympicum Larniurn garganicum Senecio rupestris Heracleum * verticillatum Scrophularia * laciniata

OTHER PHANEROGAMS

Teucrium chamaedrys Campanula velebitica Festuca sp. Melica ciliata Asyneurna canescens Cuscuta sp. Euphorbia cyparissias Urtica dioica Bupleurum sibthorpianum Cirsiurn appendiculatum Euphorbia amygdaloides Galium album Kernera saxatilis

CRYPTOGAMS

Tortula intermedia Orthotrichum cupulatum Tortella tortuosa Campanula rapunculoides Homalothecium philippeanum Schistidium apocarpum Solorina bispora

Notes: Ch - character species.

12345

EE s

sssss

EEEEE

32334

50050

34572 50000

< <

555--

77785 223.2

322.4

22233

2.22.

22 ...

.. 2 ..

.... 2

23232

22 ...

.2.32 323 ..

. 2. 2.

. 22 ..

.. 22. 32 ...

... 2.

. 2 ...

.. 2 ..

.. 2 ..

. 2 ...

253 .. . 22 ..

23 ... .. 3 ..

3 ....

3 ....

. 2 ...

e

%

100

80 80

100

60

40

20

20

100

60

60 60

40

40

40

40

20

20

20

20 20

60

40

40 20

20

20

20

folii). Bo�caiu (1971) described the Geranietum macrorrhizi from limesone screes

with a granite admixture from the Cernei Mts. (the Southern Carpathians). The altitudinal range of the latter community is, however, shifte d more to lower altitudes (between 380 to 1400 m) which suggests a classification of the Geranietum macrorrhizi Bo�caiu 1971 into the Peltarion alliaceae Horvatié 1957 (I'hlaspietea rotundifolii). Only 5 species are common both to the Geranietum macrorrhizi and Bromo-Geranietum macrorrhizi (besides the aponymous species, also Moehringia pendula,Melica ciliata, Senecio rupestris and Urtica dioica). Blecié (1958) described the Corydalo-Geranietum macrorrhizi from Montenegro (the Piva Valley in the

35

Durmitor Mts.), but only Melica ciliata and Geranium macrorrhizum are shared by the compared communities.

Syntaxonomy of the Thlaspietea rotundifolii communities

The discussed communities apparently belong to 2 alliances having their distributional optima in the alpine and subalpine belts, respectively. The Bromo lacmonices - Geranietum belongs to the Silenion marginatae Lakusié 1970 described for the calcareous-scree communities of the southern Dinarides by Lakusiè (1970) and Lakusiè et al. (1982) as suggested by lists of diagnostic species. The Silenion marginatae includes also another scree community described from the Pirin Planina Mts. - the Kenthranthetum kellereri ('I elchev & V asiliev 19 7 O). This community was reported from the altitudes between 1700 and 1900 m at one locality below the Duninoto Kuche Mt. It is similar to the Bromo-Geranietum macrorrhizi by the occurrence of Teucrium chamaedrys, Lamium garganicum, H eracleum sphondylium subsp. verticillatum and Scrophularia heterophylla subsp. laciniata. The dominant and character species of the community is Kentranthus kellereri, a Bulgarian endemie plant.

The Silenion marginatae is probably a geographic vicariant unit to the Peltarion

alliaceae distributed in the northern Dinarides and Southern Carpathians (see also above).

The alpine sere e communities show a great degree of resemblance and should be classified within one upper-ranked unit, the Veronico-Papaverion degenii all. nova hoc loco (nomenclatura! type: Papaveri degenii-Armerietum alpinae Mucina et al. 1990). The character taxa of the Veronico-Papaverion degenii comprise Veronica saturejoides subsp. kellereri, Poa pirinica, Galiùm stojanovii, Alyssum cuneifolium

subsp .pirinicum, Erigeron vichrenensis andArabis ferdinandi-coburgi as well as the

character species of the Papaveri -Armerietum and Veronico-Silenetum (see Tab. 4). This alliance, endemie to Bulgarian mountains (Pirin, probably also Rila and Slavyanka) is a vicariant to the Saxifragion prenjae and Bunion alpini described by Lakusié (1970) from the Dinarides. The synoptic table of the Dinaride scree communities (Lakusié 1970: Tab. 10a) supports the separate syntaxonomic position of the Veronico-Papaverion degenii. The discussed units are floristically different as they contain many endemie species characteristic of respective distribution areas of the alliances. The Bunion alpini, Saxifragion prenjae, Silenion marginatae and Veronico-Papaverion degenii belong to the Arabidetalia alpinaeflavescentis Lakusié 1970 (Thlaspietea rotundifolii) which is a geographic vicariant

to the Thlaspietalia rotundifolii of the Alpic-Carpathian Mountain System.

3.3. Plant Communities of Snow-bed Vegetation

The snow-patch (snow-bed) vegetation was studied both on granites, schists and

marbles in the central and nothernmost parts of the Northern Pirin Planina Mts. Two associations belonging to the Salicion herbaceae, such as the Soldanella

36

pusillae-Plantaginetum gentianoidis and Omalotheco -Alopecuretum gerardii were described from siliceous bedrocks. The Gentiano-Plantaginetum atratae and Bartsio-Salicetum reticulatae were found on calcium-rich bedrock; these latter belong to the Salicion retuso reticulatae.

Soldanello pusillae-Plantaginetum gentianoidis Boticaiu 1971 Nomenclatural type: Boticaiu (1971; Tab. 12, relevé 3), lectotypus

The Soldanello-Plantaginetum gentianoidis is a snow-patch plant community supported by siliceous bedrocks. It was noted in the southern part of the Northern Pirin Planina Mts. in the Smirnenski and Belemeto Tsirkus. The habitats of the community are found in concave relief forms; these are shallow depressions hidden among large boulders, as a rule. The snow cover persists longer than in habitats of the Omalotheco-Alopecuretum gerardii.

The shallow alpine tangel soils ,rich in fine-grained soil material are derived from granite. The soil has higher moisture retention than that supporting the Omalotheco-Alopecuretum gerardii . This is indicated also by the occurrence of Nardus stricta and Cerastium cerastoides as well as high cover values of Arenaria biflora, Taraxacum nigricans agg. and Carex curvula. Plantago gentianoides is a good moisture indicator itself. In comparison to the drier Omalotheco-Alopecuretum gerardii, the abundance of drought-tolerant taxa such as Omalotheca supina, Alopecurus gerardii, Dianthus microlepis and Scleranthus perennis subsp. marginatus decreases.

The stands are composed of two layers, including the low-grown herb layer dominated by Plantago gentianoides and subdominated by Arenaria biflora, Nardus stricta, Taraxacum nigricans agg. and Carex curvula. Mosses and lichens are represented by dominant Polytrichum piliferum; Lepraria incana, Stereocaulon alpinum and Cetraria islandica are also frequent, although theiF cover values are lower than with the Omalotheco-Alopecuretum gerardii.

Plantago gentianoides and Cerastium cerastoides are regional character species of the Soldanello-Plantaginetum gentianoidis in the Pirin Planina Mts.

As seen from Tab. 7 the Bulgarian Plantago gentianoides community can be identified with the Soldanello-Plantaginetum gentianoidis Boticaiu 1971. The floristical differences within the Soldanello-Plantaginetum gentianoidis are depicted by geographical races. The community described from the 'J;'arcu, Godeanu and Cernei Mts. (Boticaiu 1971), Fà'gà'rati Mts. (Csuros 1957, PuticaruSoroceanu et al. 1977) and the Retezat Mts. (Csuros et al. 1956) is the Luzula alpino-pilosa race, characteristic of the Rumanian Carpathians (l'ab. 7 hoc loco, columns 1-3). It is differentiated by Luzula alpino-pilosa, Festuca supina, Polytrichum noruegicum, Carex pyrenaica, Rhododendron myrtifolium (syn. R. kotchyi) and Phyteuma confusum. The Soldanello-Plantaginetum gentianoidis from the Pirin Planina Mts. is assigned to the Alopecurus gerardii race (l'ab. 7, column 5). It is differentiated by Alopecurus gerardii, Carex curvula, Achillea clusiana, Scleranthus perennis subsp. marginatus and Jasione bulgarica.

37

The Nardo-Plantaginetum gentianoidis (Lakusié et al. 1979) from the Vranica Mts. (Yugoslavia) is identical with the Soldanello-Plantaginetum. Based on taxa such as Crepis aurea subsp. glabrescens (incl. C. bosniaca Maly), Carex curta, Festuca picta and Sedum alpestre (incl. S. horakii), an eastern - Dinari de race (Tab. 7, column 4) within the Soldanella - Plantaginetum can be recognized. The homonymous "AssociationNardus stricta - Plantago gentianoides "(Ganchev 1963) from the Rila Planina Mts. is a transition-mire community found at peaty margins of glacier lakes.

Omalotheco-Alopecuretum gerardii ass. nova Nomenclatural type: Tab. 6, relevé 11, hoc loco

The Omalotheco-Alopecuretum gerardii is a widely distributed snow-bed community of the Pirin Planina Mts. It was found in broad glacial valleys and circles of the granitic Pirin Planina Mts. (the Solishcheto Saddle, Smirnenski and Belemeto Tsirkus, the Tipits Mts.). The community was sampled also in the Rila Planina Mts. as documented by the following relevé:

Rila Planina Mts., a saddle near the Vazela Mt., 2600 m, aspect ENE, slope 35°, area 16 m2, cover of

herb layer: 75%, cover of moss layer: < 1 %; August 11, 1978.

Alopecurus gerardii 3, Omalotheca supina 2-3, Campanula alpina subsp. orbelica 2 ,Dianthus microlepis

1, Poa media 1, Sedum alpestre 1, Taraxacum sp. 1, Luzula spicata+, Scleranthus perennis subsp. marginatus +, Arenaria biflora +, Polytrichum sp. +, Cetraria islandica +.

References to snow-bed communities with Alopecurus gerardii from the Rila Planina Mts. were made by Rusakova (1977) and Rusakova-Anastasova (1983). Grebenshchikov (1965) and Jovanovié et al. (1975) mentioned the occurrence of these communities also from other south-Balkan mountains.

Unlike the Soldanello-Plantaginetum gentianoidis, this community prefers convex forms of relief. The sites face various aspects; the slopes have a gentle inclination, but are still well-insolated and exposed to winds. Especially in saddles, cryoturbation forms a great variety of habitats suitable for the community. In some places the development of deeper soils is prevented by co-action of wind and snow cover, and large so il-free patches are formed. Thé bottom of the patches is covered by denuded granite boulders and gravel-like eroded material derived form the granitic rocks. Thick rhizomes of Alopecurus gerardii penetrate to the ground surface to form a unique worm -like plexus solidifying the substratum. Among these, abundant Omalotheca supina, Dianthus microlepis, Campanula alpina subsp. orbelica andScleranthus perennis subsp.marginatus occur. The lower herb sublayer is only few cm high. The culms of Alopecurus gerardii, Luzula spicata and flowering shoots of Achillea clusiana attain the maximal height of 1 O to 15 cm, and form the upper herb sublayer. The cryptogamic layer covers usually a quarter of the surface, and is composed of few mosses (the abundant one is Polytrichum alpinum) and lichens, which dominate the stands in the driest habitats (the euphrasietosum minimae).

38

Most of the stands are grazed by sheep and sustain heavy trampling as becomes o bvious along touristic tracks.

The character species of the association isAlopecurus gerardii. The group of theassociation differential species includes drought-tolerant taxa characteristic of acid grasslands of the Seslerion comosae, such as Festuca riloensis, Thymus balcanus, Campanula alpina subsp. orbelica, Vaccinium gaultheroides, Euphrasia minima,

Festuca supina andAgrostis rupestris. Also a lichen - Baeomyces roseus - should be listed among the differential species.

Three subassociations were distinguished within the Omalotheco -Alopecuretum gerardii such as the plantaginetosum gentianoidis, the typicum, and the euphrasietosum rninimae. The differentiation of the subassociations is underlied by soilmoisture gradient, and is stress ed by the degree of drought tolerance of differential species groups.

The Omalotheco-Alopecuretum plantaginetosum gentianoidis subass. nova (nomenclatura! type: relevé 6 in Tab. 6 hoc loco) is a transitional unit to the Soldanello-Plantaginetum gentianoidis. It occurs in the Belemeto Tsirkus and on the north-facing slope of the Mozgoviskhi Chukar Mt. (2605 m). The stands on gentle slopes and the soils are the richest in fine materiai within the range of the association. The community houses a number of moisture -loving species which occur only with low abundance values in the other subassociations of the Omalotheco-Alopecuretum gerardii. It is the species-poorest snow-bed community. Plantago gentianoides, Alopecurus gerardii, Omalotheca supina and Taraxacum nigricans agg. are the dominants of the Omalotheco-Alopecuretum gerardii plantaginetosum. The differential species of the subassociation are Plantago gentianoides çind Ligusticum mutellina. Taraxacum nigricans agg. has the highest cover values Ìn this subassociation.

The subassociation typicum subass. nova (the nomenclatura! type is identica! with that of the association) is a widely distributed vegetation type occupying an intennediate position within the Omalotheco-Alopecuretum gerardii along gradients of soil-moisture and content of fine-soil particles. The group of differential species of the subassociation consist of Acinos alpinus, Luzula spicata andLophozia ventricosa.

The subassociation euphrasietosum minimae su bass. nova (nomenclatura! type: relevé 18 in Tab. 6 hoc loco) occupies the driest positions along the soil-moisture gradient within the associati on. It was located in the Demirkapiiska Reka Valley and at the Tipits Mt. The stands of this community are extremely low-grown (2-5 cm); the cryptogamic layer is rich in lichens such as Lepraria incana, Stereocaulon alpinum, Cetraria islandica, Cladonia macrophyllodes and Solorina crocea. The differential species of the subassociation include Euphrasia minima, Agrostis rupestris, Festuca supina, Ranunculus pseudomontanus, Potentilla aurea subsp. chrysocraspeda, Nardus stricta, Plantago atrata and Hypnum cupressiforme and Rhacomitrium canescens. The Omalotheco-Alopecuretum euphrasietosum is a snowbed community of siliceous substrata, with a high abundance of Plantago atrata which, however, is more frequent in snow-beds on calcareous bedrocks. Incidental-

39

Tab. 6 - Snow bed communities on siliceous substrata.

No. of relevé

Exposition

Slope 0

Sampled area m 2



CARICETEA CURVULAE

Dianthus microlepis Carex curvula Poa media Scleranthus * marginatus Primula minima Jasione * bulgari ca Juncus trifidus Poa laxa

111111111122 1234 56789012345678901

W E N N S N SS

WWNWWWWSESWWNW-WN

1 11<1215<1 11 12 55005050050505-50

1 11 1 1 1 8904 22933342608928649

1111 1 9997 00008669795678906 0050 00000000000050000

<22 2 2 11 <33321< 2 1001 0550-555500000555

. . 3. .. 2322.352223453 . 7343 32.2.23 .. 2.53 ... 3 2.35 .. 4.22 .... 553 .. 42 .12. .... 2 .. 52 ... 323.2

3 ................ .. 4. .... 2 ......... 3.

""" .3. · '·

LIGUSTICO-PLANTAGINETUM GENTIANOIDIS

Cl C2 % %

25 76 100 53

75 47 50 41

o 6 25 12

o 6 o 6

Plantago gentianoides (Ch)

Cerastiurn cerastoides (Ch) 8887 778 ...... 2...... 100 24 2 .. 2 . . . . . . . . . . . . . . . . . 50 O

OMALOTHECO-ALOPECURETUM GERARDII

Alopecurus gerardii (Ch) Campanula * orbelica (D) Geum montanum (D) Festuca riloensis (D) Thymus * polytrichus (Dl Baeomyces roseus (D) Vaccinium gaultheroides (D)

.2.3 77778777777887777 2.22223322.223246 2 ....... 1.22 ... 12 ... 2.323 ........ 5 .... 322.2 ...... 2. 4 ... :.222 ....... . 2 ..... 2 .. 2 ...... .

D-EUPHRASIETOSUM MINIMAE

Euphrasia minima ............. 4334 Agrostis rupestris ............. 3223 Festuca supina ............. 456. Rhacomitrium canescens ............. 4 .. 2 Ranunculus pseudomontanus . . 3. 22 .... 21.2.2.555 . Nardus stricta 3743 .. 2 .... 2 .. 553. Potentilla * chrysaspeda 3. 4. ... 2 .. 22 ..... 343. Plantago atrata . . 3. . · .. 2 ......... 442 . Hypnum cupressiforme . 2 .. ............. 242 .

SALICETEA HERBACEAE

Omalotheca supina 2323 73375557374778767 Arenaria biflora 7474 .2223 .... 2323 .. 2. Achillea clusiana 2 .. 2 ... 5 .... 32322 .... Sedum alpestre ... 2 ....... 1222.3 .. 22 Ligusticum mutellina 32 .. .22 .............. Luzula spicata ..... 232 ......... Soldanella pusilla ... 3 .......... 2 ...... Ranunculus crenatus ".2 "" ..... 1 ....... Luzula spadicea ................ 2

OTHER PHANEROGAMS

Taraxacum sp. 7237 553.2 ... 12333 .... Acinos alpinus ... 22. 2 .......... Jovibarba heuffelii ........ 1. .......

40

50 100 O 88 O 35 O 29 O 29 O 24 O 18

o 24 o 24 o 18 o 12

25 53 100 29

50 35 25 24 25 18

100 100 100 53

50 35 25 41 50 12

o 18 25 6 25 6

o 6

100 53 o 18 o 6

Thymus X pilisiensis . . . . ... 3 ............. o 6

OTHER CRYPTOGAMS

Polytrichum piliferum 2752 2245.33 .. 67744444 100 82

Cetraria islandica 2 ... 2 .. 2.222 ... 222222 25 65

Lepraria incana . 34. .. 2 ....... 3345236 50 47

Stereocaulon alpinum . 22. .2 ... 23 ... 2 .. 22 .. 50 35

Lophozia ventricosus ... 2 .......... 345 .... 25 18

Cladonia rnacrophyllodes . . 2. ............ 22.2 . 25 18

Solorina crocea ............ 2 ... 2 o 12

Cladonia pyxidata ...... 2 .......... o 6

Cladonia sp. ................ 2 25 18

Lecidea sp. ....... 2 ......... o 6

Myurella julacea 2 .... ............. 3 ... 25 6

Notes: X - hybrid; for other abbreviations see Tab. 1.

ly, this is the only calciphilous species which does occur on silicate snow-beds. On the other hand, the number of the acidophilous elements growing also on calcareous bedrocks is much higher · (eg. Gentiano -Plantaginetum atratae trifolietosum orbelici; Tab. 7).

Simon (1958) considered Alopecurus gerardii to be a character species of the Seslerietalia comosae. Horvat (in Horvat et al. 197 4) described the AlopecuroPlantaginetum from Pelister Mts. (Yugoslavia Macedonia). According to Horvat et al. G.c.) this community has an outrageous position within the Seslerion comosae as it occurs on fine-grained soil and at less exposed habitats then other communities of the alliance. It differs from the Omalotheco-Alopecuretum by presence of the association character species such as Plantago holosteum, Lotus corniculatus and Thesium alpinum, and by absence of many species of the snow-bed habitats. Alopecurus gerardii (var. pantocsekii) is one of the dominating species of the Ranunculetum crenati Lakusié 1966 described from Bjelasica Mts. in Yugoslavia.

Quézel (1964, 1967) regardedAlopecurus gerardii as a character species of the Caricetea curuulae. Three associations dominateci by the species were described by Quézel G .c.) from the Greece mountains, belonging to the Trifolion parnassi Quézel 1964 (the Trifolietalia parnassi Quézel 1964, Caricetea curuulae). The community group is vicariant to the Nardion Luquet 1926 of the temperate Europe. The communities are confined to shallow hollows or small-sized dolinas filled by silt. These habitats contain snow longer than the surrounding environments although they cannot be classified as true snow-beds characteristic of the temperate and boreal mountain ranges. In summer, the soil surface in the sites desiccates. The Croco sieberi -Alopecuretum gerardii Quézel 1964 (syn. Beta nanae-Alopecuretum gerardii (Quézel 1964)Horvat et al. 197 4) was described from the Taygetos, Kyllini and Parnassos Mts., the Gnaphalio hoppeani-Alopecuretum gerardii Quézel 1967 is known from the Thessalian Olymbos, and the Croco ueluchensis-Alopecuretum gerardii Quézel 1967 was found to occur in the central and northern parts of the Pindos Mts. (Quézel 1964, 1967, Horvat et al. 1974).

AnalogousAlopecurus gerardii communities were described from the Djurdijura Mts. in north Africa (Quézel 1957), Maritime Alps in France (eg. Guinochet 1938, Barbero 1970, Lacoste 1975: Ranunculo pyrenei - Alopecuretum gerardii) and Pyrenées (Ludi 1943, Braun-Blanquet 1948: Trifolio - Phleetum gerardii).

41

Tab. 7 - Communities of the Salicetea herbaceae (with Plantago gentianoides. Ranunculus crenatus and Alopecurus gerardii) in the Balkan Peninsula.

Number of column 10 11 12 13

Relevés per table 10 10 17 ---------------------------------------------------------------------------------------------------------------

SALICETEA HERBACEAE

Plantago gentianoides V24 V+3 V15 434 434 V+ II23 II+ III+ II+4 Ranunculus crenatus I+ II+l II+ l+ 1+ V23 V24 423 Vl3 V24 V+3 V14 Ir Alopecurus gerardii 2+1 III13 II12 V13 V34

Omalotheca supina III+ III+ IIIr+ 21 4+1 V+2 V+l 2+3 IV+2 II+ V15 V14 v2, Geum montanum IV+ II+ III+l 2+1 V+2 III+ 11 II+ V+l V+2 V+2 IIr+ Luzula alpino-pilosa V+l I+ IIl III+ V+l 2+ Il+l IV+2 I+ Sedum alpestre III+ l+ V+l II+ 11 I+ I• III+2 IV12 IIIrl Soldanella pusilla V+2 V13 V+l 11 4+2 V14 I+ Ligusticum mutellina V+2 III+ V13 412 2+1 V23 IV+2 4+3 II+ IV+l IV+2 I+ Polytricum norvegicum I+ III+l IV+2 I+ II+ Arenaria biflora IV+ 423 2+2 I2 III+l Tanacetum alpinum I+ I+ III12 IV+ I2 Cerastium cerastoides I+ 2+ IIl IV+2

Plantago atrata 11 I+ Il3 !Il) IT+2 Achillea clusiana 2+ JI+ 1 Luzula spicata J+ I+ Salix herbacea Il I+ Kiaeria starkei III+J I4 Soldanella hungarica V+2 Kiaeria falcata III23 Anthelia juratzkana III+2 Polygonum vi viparum I+ Veronica alpina Il

CARICETEA CURVULAE

Potentilla chryocrasped;i IV+l I+ III+2 31 212 V+2 12 II+ IV+ IV+2 Vl3 II12 Festuca supina Il III+5 IV+ 1+ II+ I+ I2 Primula minima I+ V+2 IV+l l+ III+l IV+l rl Agrostis rupestris V12 I+ II12 I+ Il II+1 Juncus trifidus II+ II+ I+ II+ Il Carex curvula 413 II+l 12 I+ Poa media 11 3+3 Il2 III+3 Ranunculus montanus agg. 11 I+ lI+l Illr2 Campanula alpina II+ I+ II+ Hieracium alpinum I+ II+ I+ Pulsatilla alba II+ II+ I+ Phyteuma confusum Il+l II+ I+l Scleranthus . marq] natus 2r+ IIl III+3 Festuca riloensis V+2 11+2 Jasione orbiculata 111+2 V+<

Anthoxanthum alpinum 11 I+ Leontodon helveticus 11 Il Dianthm11 microlepis 11 III+2 Jasione bulo-arica 12 I+l Poa disparilis Il III+2 Campanula tatrae III+ Campanula . orbelica V+2

A.venula versicolor I+

Gentiana punctata I+ Oreochloa disticha I+ Anthemis carpatica l+ Festuca scardica II12

Jovibarba heuffelii Ir

Euphrasia minima 1112


Cardami ne rivularis l+ Saxi fraga muscosa I+ Doroni cum carpaticum 1+ Poa laxa Il Crepis glabrescens 31 II13 II14 Myosotis alpestris I+ Festuca picta 412 Cardaminopsis ovirensis I+ Arabis alpina I+ Saxifraga androsacea I+ Il

OTHER PHANEROt.AMS

Milrdus stricta V+2 Il 434 413 I+ IIl Vl2 II+2 I+ II+2 Taraxacum sp. div. II+ 4+3 V+ 2+1 I+ II+ Il T+ Poa alpina I+ IV+2 II+ 2+2 III+ III+l IV+l 11+1 Homogyne alpina V+l 21 V+ TV+l Il

Oeschampsia caespi tosa 11 I+ II+l Il Phleum alpinum I+ l+ TV+2

42

Carex pyrenaica II13 I+ I+

Alchemilla glaucescens I+ 2+1

Veratrum album I+ I+ I+

Soldanella alpina 2+ Il

Avenella flexuosa 1+ Il

Luzula sudetica 11 Il

Carex foetida II13 II14

Crocus heuffelianus II+

Saxifraga heucherifol ia I+

Trifolium pallescens I+

Carex e urta 3+1

Poa sp. 11

Gentiana acaulis 1+

Crocus neapoli tanus 11 Euphrasia stricta IV+

Achillea distans I+

Alchemilla glabra I+

Saxifraga stellaris I+

sesleria bielzii II+

Primula veris I+

Homogyne discolor Il

Alchemilla plicatula Il

Si lene pusilla Il

Luzula campestris I+

Liousticum albanicum III+l

Thymus . pilisiensis Il

Thymus . polytrichus II+l

CETRARIO-LOISELEURIETEA

Cetrarié1 islandica II+ I+ 1+ I+ IV+l II12 IV+

Vaccinium gaul theroides I+ I+ I+

Vaccinium myrtillus I+ 1+

Rhododendron myrtifolium II+l I+

OTHER CRYPTOGAHS

Polytrichum piliferum 11 4+3 I+ V+)

Polytrichum juniperinum 11 II+ V+3

Dicranum scoparium II+ II+

Lepraria incana 212 1II+2

Stereocaulon alpinum 2+ II+l

Cl adonia macrophyllodes l+ I+

Hyurella julacea 1+ Il

Lophozia ventricosa I2 Il2

Hypnum cupres si forme 1+ I+2

San ionia uncinata I+ I+

Polytrichum alpinum I+

Oligotrichum hercynium 1123

Diplophyllum taxifol ium II+l Dicranella heteromalla II+l Cl adonia pyxidata I+

Tortella tortuosa Il

Rhacomitrium canescens T+2

Solorina crocea I+

Cl adonia sp. I+

Baeomyces roseus 11+2

Localities of tables (Tab. 7):

1. Soldanello (pusillae)-Plantaginetum gentianoidis; Resmerità' (1976), Tab. 8; Rodna (Rumania)

2. Soldanello pusillae-Plantaginetum gentianoidis; Resmerità' (1979), Tab. 6. Rodna (Rumania)

3. Ligustico-Plantaginetum gentianoidis; Mucina (ined.); Fà'gà'ra4 (Rumania)

5. Ligustico-Plantaginetum gentianoidis; Tab. 6 in this study, relevés 1-4; Pirin

6. Agrosteto (alpinae)-Ranunculetum crenati; Resmerità' (1975), Tab. 2; Maramure4 (Rumania)

7. Soldanello hungaricae-Ranunculetum crenati; Coldea (1985); Tab. l; Rodna (Rumania)

8. Soldanello pusillae-Ranunculetum crenati; Mucina (ined.); Fà'gà'ra4 (Rumania)

9. Soldanello (pusillae) - Ranunculetum crenati; Bo4caiu (1971), Tab. 24, '{'arcu, Godeanu & Cernei Mts. (Rumania)

10. Soldanello (pusillae)-Ranunculetum-crenati; Resmerità' (1976), Tab. 3; Retezat Ms. (Rumania)

11. Ranunculetum crenati vranicensis; Lakusié et al. (1979), Tab. I (7 rels.); Vranica (Yugoslavia)

12. Ranunculetum crenati; Lakusié (1964-1966), Tab. 2, relevés 1-2, 6-12; Bjelilo (Yugoslavia)

13. Omalotheco-Alopecuretum gerardii; Tab. 6 in this study, relevés 5-21; Pirin

43

Gentiano-Plantaginetum atratae ass. nova Nomenclatura! type: Tab. 8, relevé 9, hoc loco

The Gentiano-Plantaginetum atratae is a typical snow-bed community of the northern range of the Pirin Planina Mts. It is limited to habitats which are covered app. 6 months by snow and occur at altitudes between 24 00 and 2600 m. The community is found, unlike the Bartsio-Salicetum reticulatae, on convex relief forms, usually with gentle (10 to 15°), east to south - (southeast) -facing slopes. The habitats are more insolated but also windier than those of the Bartsio -Salicetum reticulatae.

The soils, shallow alpine rendzinas, are derived from two types of rocks such as calcium-rich schists and crystalline limestones (marble). The substratum is very skeletal; the skeleton covers a small part of the soil surface. On schists, the upper soil layer is rich in gravel-like skeleton.

The stands of the community and low-grown and two-layered. The herb layer, usually with high cover ( 70 to 90%) is composed of two sublayer which are not always developed. The low sublayer (3 to 5 cm) is dominateci by Plantago atrata which gives the stands an outlook of a silvery carpet. Only occasionally some grasses and sedges, such as Alopecurus gerardii, Poa pirinica, Sesleria coerulans and Carex kitaibeliana overtop the sublayer and form the higher herb sublayer usually attaining l0to 15 cm in height. The dense carpet of Plantago atrata is penetrateci by dwarf herbs such as Gentiana uerna, Ranunculus carinthiacus (see Kozuharov & Petrova 198 8), Omalotheca supina, Arenaria biflora, Sedum atratum, Potentilla crantzii, P. aurea subsp. chrysocraspeda, Dianthus microlepis, Campanula alpina subsp. orbelica, Euphrasia minima, Galium stojanouii, Oxytropis urumouii and Draba scardica.

The community is found in patches aminds of calcareous alpine grasslands. The appearance of stands of this community indicates a good potential to sustain trampling as we also observed in areas heavily affected by mountain walkers and alpinists, for intance in the Premkata Saddle (2650m). The stands of the Gentiano -Plantaginetum have not changed markedly in this location during the observed period of 1978-1984, although the number of tourists kept increasing year to year. The dense low-grown carpet serves a good anti-erosion function.

The character taxa of the Gentiano -Plantaginetum are Plantago atrata, Gentiana

uerna, Potentilla crantzii and Ranunculus carinthiacus. The group of differential species of the Gentiano-Plantaginetum is composed of a number of indicators of siliceous substrata, such as Alopecurus gerardii, Dianthus microlepis, Campanula alpina subsp. orbelica, Acinos alpinus, Euphrasia minima, Trifolium repens subsp. orbelicum and the like (Tab. 8). Asplenium fissum and Alyssum cuneifolium su bsp. pirinicum, which are characteristic of skeletal soils or rocky habitats, can be also considered differential for the Gentiano-Plantaginetum. Of cryptogams Leskea polycarpa, Psora decipiens and P. lurida are also differential for the GentianoPlantaginetum.

Two subassociations were distinguished within the association, the typicum

44

Tab. 8 - Communities of the Salicion retusae.

No. of relevé 1111111

123456789 0123456

Exposition ss E

N SS N N N

ESEEEESEW EN-ENW-

Slope 0 1111221 1 23 223<

500005050 00-0005

Sampled area m2 1111111 111 042652289 4750226

Cover of herb layer % 1

999789799 0899887

055050005 0000505

Cover of mass layer % 1<21< <1 <1< 1 6

05051-150 1055050

GENTIANO-PLANTAGINETUM ATRATAE

Plantago atrata IChl 888778888 . 2. 32 ..

Gentiana verna IChl 244433323 . 3. 22 ..

Acinos alpinus IDl 22.5.4.2.

Leskea polycarpa (Dl ... 3 .. 22.

D-TRIFOLIETOSUM ORBELICI

Campanula * orbelica 222.23 ...

Oraba scardica .12222 ... Trifolium . orbelicum . 362. 5 ...

Alopecurus gerardii .. 6346 ...

Euphrasia minima . 352.

Botrychium lunaria . 2. 4 . ....

Dianthus microlepis 222.3. . .... 2.

Artemisia eriantha . 3. 6. 2 ... . 1. .. 3.

Potentilla * chrysaspeda .. 55. . . 23 ...

BARTSIO-SALICETUM RETICULATAE

Salix reticulata !Chi 9899848

Bartsia alpina (Ch) . 22. 43.

Polygonum viviparum ICh) 33.52.2

Erigeron vichrenensis IChl .. 2. .122 .. 2

Dryas octopetala (Chl ... 7.

Di trichum flexicaule IDl .532335

Hnium stellare ID) . 3 .. 322

Pinguicula balcanica IDl . 2 .. 52.

Saxifraga androsacea IDl . 3 .... 2

Saxifraga ferdinandi-coburgi IDl 2 .... 2.

Silene * graefferi IDl 2 .. 2 ...

Saxifraga oppositifolia IDl ... 2 . . . 2. .42.335

Primula minima IDl .. 2 .. .... 5. 5

ARABIDETALIA CAERULEAE

Carex kitaibeliana . 525.2.32 .32557 .

Carex • pirinica .. 22 ... 56 .3323.2

Poa pirinica .. 5.2342. 3. 3 ... 2

Potentilla crantzii 364444.37 3.23.

Ranunculus carinthiacus . 2332 .. 53 3 .. 3 .

Sedum atratum 22122. .22.

Primula halleri . . . 2 . . . 22 3111

Viola grisebachiana . . . 2. 23 .

SALICETALIA HERBACEAE & SALICETEA HERBACEAE

Omalotheca supina Arenaria biflora Phleum • rhaeticum Veronica alpina

Achillea clusiana Geum montanum Sedum alpestre Luzula spadicea

45

.22422253 222.

2322.2355 2222 .. 2

...... 22 ... 2 ..

.... 2 .. 2 ... 2

... 3 ...

.. 2.

... 2 ..

.... 2.

Cl

%

100

100

56

33

56

56

44 44

33

22

44 33

22

o

o

o

11

o

o

o

o

o

o

o

22

11

67

44

56

89

67

56

33

11

89

89

22

11

11

11

11

11

C2

%

43 43

o

o

o

o

o

o

o

o

14 29

29

100

57

71

57

14 86

57

43 29

29

29

71

29

71

71

43

43 29

29 57

29

43 71

14 29

o

o

o

o

ARABIDETALIA ALPINAE-FLAVESCENTIS

Armeria * alpina Galium stojanovii Myosotis alpestris Veronica * kellereri Thlaspi * bellidifolium Saxifraga * discolor Alyssum * pirinicum Arenaria pirinica Saxifraga luteoviridis Arabis caucasica Myosotis suaveolens


Doronicum columnae Polystichum lonchitis Sagina saginoides Geum reptans

FESTUCO-SESLERIETEA

Cerastium * lanatum Sesleria coerulans Pedicularis verticillata Poa alpina Oxytropis urumovii Androsace * arachnoidea Anthyllis * vitellina Aster * dolomiticus Onobrychis scardica Trifolium badium Carex rupestris

CARICETEA CURVULAE

Saxifraga * exarata Scleranthus * marginatus Festuca riloensis Hieracium alpicola Leontodon * riloensis Poa ursina Sesleria comosa Knautia midzorensis Juniperus * nana


Silene pusilla Asplenium fissum Euphrasia salisburgensis Minuartia verna Potentilla * stojanovii Thymus cherlerioides

OTHER PHANEROGAMS

Taraxacum nigricans agg. Gentianella sp.

OTHER CRYPTOGAMS

Tortella tortuosa Hymenostylium recurvirostre Cladonia sp. Polytrichum juniperinum Bryum caespiticium Psora decipiens Psora lurida Catopyrenium cinereum Cladonia symphycarpa Tortulla sinensis Trichostomum crispulum Bryum elegans Cetraria islandica Cladonia pocillum Peltigera rufescens Preissia quadrata

46

.2252.352 24252325. .. 22.2322 . 2. 33. 3 .. 22. 2 .. 2 ..

.. 2 .1. . ...... 22.

... 23.

.. 2 .....

..... 2.

24 .. 323 2322 ... 32. 2 .. 2 232 ... 2 .2 .... .

.. 2

.. J.

..... 2

.... 3. .2 .. 1.2

.... 2. ....... 2 .2.

.. 5.

75323.J. 22 .2323.42. ... 2 ... 2. . 3. 2 ... 63 . 5 .. 3. ..... 2. 3. 2.

,,], .. 3. .. .. 6.

... 2. 2. 2. 2 ... .. 4. 2 ...... 2 .. ... 2 ...

3. 22 .. 3 433443. 22.332. . 235 .. 2 .. 2. 2. ... 2.

.1 .. 2.

33 .

... 3

.. 1 ....

... 2 ..... . 2 .... 3

... 2 .. 2 .. .2 ....

2 ... .. 1.

. 2.

.2.23.432 232.2 ..

. . 32 .. 2. 2 . 5 .... 5

332 .. .. . 35. 3 ... . .. . 2 .. .

.2 ...... 2

... 3 ... 2.

.2 .. .. ..... 3

• 2 •. ...... 2 ..

... 2 ...

22.2.5 . .52.53 . ... 2. ... 5 .. . .2 .... .

2 ... 2 .. ..... 23 ..... 37 .. 2 ... 3 .2 .... 2

78 89 67 44

44

22 22 22 11 11

o

11 11 11

o

89 67 22 44 22 11 11 11 11 11 11

11 11 11 11 11 11 11 11

o

11 22 11 11

o 11

67 o

44 22 33 33 11 22 22 11 11 11 11

o o o o o

71 57 57 57 14 14

o o

14 o

14

43 o

14 14

57 86 71 43 29 14 14 14 14

o 29

14 o o o o o o o

14

29 o o o

14 o

57 14

57 57 14 14 14

o o o o o o

29 29 29 29 29

Solorina bispora Blepharostoma trichophyllum Collema sp. Crataneuron filicinum Distichum capillaceum Hypnum cupressiforme Lepraria incana Lophocolea minor Stereocaulon alpinum

. 2 ... 3. ..... 2.

. 2 ..

. 3 .....

2 ....•.

... 3 ...

..... 2.

..... 2

...... 2

O 29 O 14

O 14

O 14

O 14

O 14

O 14

O 14

O 14

Notes: Cl - constancy of Gentiano-Plantaginetum: C2 - constancy of Bartsio-Salicetum; far other abbreviations see Tab. 1.

subass. nova (the nomenclatura! type is identica! with that of the association) and the trifohetosum orbelici subass. nova (nomenclatura! type: relevé 3 in Tab. 8 hoc loco). The differentiation of the subassociations is based both on character of bedrock and presence of species groups. The typical subassociation occurs on marble, where as the trifolietosum orbelici is found on calcium -rich schists characteristic by plate separation of comparative more fragile bedrock. This type of bedrock is found usually along tectonic faults at contact zones between granites and crystalline limestones. The community was localized in the Kabata and Premkata Saddles, south and north of the Vikhren Mt., respectively, and near to the

Kamenishki Vrakh Mt. (2533 m). The stands were found on gentle, prevailingly east-facing slopes. The south aspect is common with the typical subassociation. The trifolietosum orbelici is a typical ecotone community housing the highest

number of vascular species of all snow-bed communities in the area regardless the bedrock. The community is a transitional unit between the Gentiano-Plantaginetum

atratae and Omalotheco-Alopecuretum gerardii. The differential truca of the trifolietosum orbe lici (I'rifolium repens subsp. orbelicum, Alopecurus gerardii, Sedum atratum, Dianthus microlepis, Campanula alpina subsp. orbelica, Potentilla aurea subsp. chrysocraspeda, Euphrasia minima) are typical of the silicate substrata (see Simon 19 5 8). Further also A c inos alpinus, Thymus cherlerioides, Artemisia eriantha, Draba scardica, Botrychium lunaria and Arenaria pirinica, which are limited to calcareous substrata, are considered differential of this subassociation. Achillea clusiana, Scleranthus perennis subsp. marginatus, Sedum alpestre, Primula minima,

Geum montanum, Festuca riloensis, Luzula alpino-pilosa, Poa media, Hieracium alpicola, Sesleria comosa and Leontodon croceus subsp. riloensis are found only in this type of snow-bed communities on calcareous bedrock, but their ecologica!

optimum lies either in snow-bed habitats on silicate bedrock or on acid alpine grasslands of the Seslerietalia comosae.

Plantago atrata includes several subspecies and varieties limited to particular mountain ranges in the Central and south Europe and forma vicariant group of taxa (Hayek 1931, Casper 197 4). Feoli-Chiapella &Feoli (1977) reported on aPlantago atrata var. tenuis dominated community from the Majella Mts. in the Central Apennines. According to the character of habitats and floristic composition (several vicariant taxa occurring in respective communities) the Gnaphalio-Plantaginetum atratae Feoli-Chiapella et Feoli 1977 is a vicariant to the Gentiano-Plantaginetum atratae.

The Trifolio-Plantaginetum angustifoliae, described by Lakusié (1966) from Bjelasica Mts. in Yugoslavia is floristically probably the most similar community to

47

the Gentiano-Plantaginetum atratae. From the Gentiano - Plantaginetum the former one differs by dominance of Crepis aurea subsp. glabrescens (syn. C. columnae), Soldanella alpina, Taraxacum spp. and Trifolium pallescens and other species of limited distribution (see Tab. 7).

Other Plantago atrata dominated communities from the Balkan Peninsula were described from silicate snow-beds. The Plantaginetum atratae studied in the Durmitor, Cvrsnica and Prenj Mts. in Yugoslavia by Horvat et al. (197 4) differs from the Gentiano-Plantaginetum atratae by presence of taxa such as Crepis aurea subsp. glabrescens, Arenaria rotundifolia, Armeria canescens and Viola calcarata subsp. zoysii and several other, mostly acidophilous, taxa. The SoldanelloPlantaginetum durmitorei, mentioned from the Durmitor Mts. without a relevé table (Lakusié 1984), is probably identica! with the Plantaginetum atratae. The Thlaspi microphylli-Plantaginetum atratae (see Horvat et al. 197 4) known from the Yugoslavian Macedonia is characterized by Androsace hedraeantha and Ranunculus crenatus. Together with the Plantaginetum atratae are, unlike the Gentiano -Plantaginetum atratae, classified within the Salicion herbaceae.

Bartsio-Salicetum reticulatae ass. nova Nomenclatura! type: Tab. 8, relevé 11,hoc loco

The Bartsio-Salicetum reticulatae is another type of a snow-bed community on calcareous substrata. The habitats of the association are exposed to longer-lasting snow period than in the case of the Gentiano-Plantaginetum atratae. The community inhabits concave forms of relief sheltered from direct solar irradiation. The stands are found in shallow depressions among large boulders on bottoms of glacial circles or on small-sized terraces with a gentle north-facing slopes.

The soils are alpine rendzinas which contain more silt than those supporting Gentiano-Plantaginetum atratae. As a result of higher contents of finer soil particles and long-lasting snow cover the soils are wetter and keep moisture even during hot summers. The desiccation is prevented also by high plant cover which attains 80 to 100% as a rule. The soils are derivedfrom crystalline limestones in all of the studied stands.

The Bartsio-Salicetum reticulatae is formed by low-grown stands composed of the herb and moss layers. The lower herb layer is dominated by Salix reticulata. Only in one relevé the dominating species was Dryas octopetala. The dominants are accompanied by dwarf alpine plants such as Bartsia alpina, Erigeron uichrenensis Pawlowski,Polygonum uiuiparum, Primula minima, P. halleri, Veronica saturejoides subsp. kellereri, Pedicularis verticillata and other. The stands of the community are recognizable as dark-green patches. The upper herb sublayer is species -poor (Carex kitaibeliana, C. parviflora subsp.pirinica, Sesleria coerulans, Poa pirinica, Armeria pocutica subsp. alpina), and does not overshoot the height of 15 cm. The moss layer is richer in species than with the Gentiano-Plantaginetum atratae stands, which also points upon the higher soil moisture. Saxifraga oppositifolia, S. androsacea, Pedicularis verticillata and Primula minima also prefer moister

48

habitats. The stands of the Bartsio-Salicetum reticulatae do not suffer frorn sheepgrazing (perhaps only from slight grazing of rnountain goats) as they occur in remote sites in bouldery glacial circles and on exposed terraces.

The character species of the Bartsio-Salicetum reticulatae are Salix reticulata,

Bartsia alpina, Dryas octopetala, Polygonum viviparum, Gentianella aspera and Erigeron vichrenensis. The latter taxon, endemie to the Pirin Planinn Mts. (Pawlowski 1969), has its sociological optimum in the Bartsio-Salicetum reticulataeand to a lesser extend it also occurs in the Papaveri-Armerietum. The other character species of the Bartsio-Salicetum are typical regional character species which occur also in other European high-mountain systerns where they show varying sociological valency. Saxifraga oppositifolia, S. androsacea, S. ferdinandi-coburgi,

Pinguicula balcanica, Silene ciliata, Primula minima, Ditrichum flexicaule, Mnium stellare, Bryum elegans, Cladonia pocillum, Peltigera rufescens, Preissia quadrata and Solorina bispora differentiate the Bartsio-Salicetum reticulatae from the Gentiano -Plantaginetum atratae.

The Bartsio-Salicetum reticulatae belongs to a group of vicariant plant associations where in also the Salicetum reticulatae frorn the West Carpathians, Salicetum retuso-reticulatae from the Alps, Dryado-Salicetum reticulatae and Salicetum retuso-kitaibelianae frorn Yugoslavia (Braun-Blanquet & Jenny 1926, Szafer et al. 1927, Horvat 1936, Beldie 1967, Lakusié 1970) are classified. The Saxifrago sempervivi-Salicetum reticulatae (Horvat 1936) appears as floristically dose to the Bartsio-Salicetum reticulatae, although the occurrence of Saxifragasempervivum, Omalotheca hoppeana, Androsace hedraeantha and Saxifraga glabella in the Saxifrago-Salicetum emphasize an important floristic difference between the discussed units. Whereas the Salicetum retuso-reticulatae, Salicetum reticula

tae and Dryado - Salicetum reticulatae belong to the Arabidion caeruleae Br.-Bl. 1926, the Saxifrago-Salicetum reticulatae, Salicetum retuso-kitaibelianae and Bartsio-Salicetum reticulatae are classified within the Salicion retusae Horvat 1949.

Syntaxonomy of the Salicetea herbaceae communities

All known snow-bed communities in Europe are classified within the Saliceteaherbaceae (Braun-Blanquet 1976, Dierssen 1984). The bedrock differentiation is reflected on the level of orders. The Salicetalia herbaceae and Arabidetaliacaeruleae comprise communities on silicate and calcareous bedrocks, respectively. A number of vicariant alliances were described within each arder (see Dierssen 1984 for a review).

The snow-bed plant communiti�s of the Balkan mountain ranges (including the Southern and Eastern Carpathians) used to be separated into the Ranunculioncrenati (Lakusié 1966, 1970), supposedly a geographic analogon to the Sali,eionherbaceae of the Alps and Carpathians (Ludi 1921,Braun-Blanquet &Jenny 1926, Krajina 1922, Riibel 1933, Oberdorfer 1977). Ranunculus crenatus, Soldanellapusilla, Plantago gentianoides were listed by Lakusié (1966) as character species of the Ranunculion crenati. Ranunculus crenatus is distributed mainly in the Balkan

49

Peninsula, but also occurs in Styria (Steiermark, Austria) in the Rottenmanner Tauern Mts. and Schladminger Tauern Mts. (Muller & Baltisberger 1984). Plantago gentianoides is a Balkan endemie, but it occurs also in the Caricion fuscae Koch 1926 em. Klika 1934 (Juhasz-Nagy 1963,Mucina ined.);Soldanella pusilla is common to many Balkan mountains, but occurs also in the Alps in the Salicion herbaceae (Braun-Blanquet & Jenny 1926, Braun-Blanquet 1954, Oberdorfer 1977).

Salix herbacea, Polytrichum noruegicum, Kiaeria starkei, K. falcata, Sibbaldia procumbens, Anthelia juratzkana and many others (Dierssen 1984), considered the character species of the Salicetea herbaceae and Salicion herbaceae, occur in many snow-bed communities in the Balkans (Horvat et al. 1937, Simon 1958, Lakusié 1966, 1970, Resmerità' 1975, 1978, Coldea 1985, Coldea et al. 1981 etc.).Horvat et al. (1937), Beldie (1967), Pu!icanu-Soroceanu et al. (1956), Popescu et al. (1983), Resmerità' (1975, 1978, 1979) and many other report also the Salicetum herbaceae Br. - Bl. 1913, Luzuletum spadiceae Br. - Bl. 1926 and Polytrichetum sexangularis from the Rumanian Carpathians and Bulgaria. However, we do not identify the Salix herbacea, Luzula alpinopilosa and Polytrichum noruegicum dominated com -

munities of the Alps (belonging to the Salicion herbaceae) with those of the Balkans. The latter should be considered vicariant units, like in the case of the WestCarpathian snow-patch communities (Dtibravcova in Mucina & Maglocky 1985).

W e do not consider the separation of the Salicion herbaceae and Ranunculion crenati for readily documented, because besides a group of snow-bed communities with their distribution areas exclusively located in the Balkan Peninsula (see the references ab ove and Tab. 7), there are some others (eg. the Poo -Cerastietum cerastoidis (Soyrinki 1954) Oberd. 1957 andNardo -Gnaphalietum supini Bartsch 1940) which occur both in the Alps as well as Southern and Eastern Carpathians (already in the Balkans).

It is a generai phenomenon that the syntaxa characteristic of calcareous bedrocks in the Balkans can be classified into more vicariant units than those from silicate rocks. This probably goes on the account of high endemism on calcium-rich

substrata, relict character of the habitats, higher species-richness, the extent of calcareous substrata especially in the Dinarides, and long-lasting isolation of the mountain summits.

Also the floristic differentiation of the high-ranked syntaxa of communities on the calcareous bedrock is more pronounced. This holds also for the differentiation of the Arabidion caeruleae (the Alps, West Carpathians) from the Salicion retusae (the Balkan mountains).

Both Gentiano-Plantaginetum atratae and Bartsio-Salicetum reticulatae are classified within the Salicion retusae, an alliance originally described from the Dinarides by Horvat (1949). Balkan endemics such as Saxifraga sedoides subsp. prenja, S. semperuiuum, S. glabella, Plantago atrata var. angusti/olia Hal. et Bald. and Androsace hedreantha are very typical of the alliance.

The SaUcion retusae belongs to the Arabidetalia coeruleae. The status of the Salicetalia retusae, a unit suggested by Lakusié (1970), still remains to be cleared.

50

The syntaxonomic relations of the syntaxa of snow-bed communities in the Balkan Peninsula can be summarized as follows:

Salicetea herbaceae Br.-Bl. 1947 Salicetalia herbaceae Br.-Bl. 1926 Salicion herbaceae Br.-Bl. 1926 (syn. Ranunculion creanti Lakusié 1966)

1. Ranunculo crenati - Salicetum herbaceae (Horvat 1936) Mucina et al. nom.novum hoc loco(basionym: Salicetum herbaceae balcanicum Horvat 1936; syn. Salicetumherbaceae sensu auct. balcan.)

2. Ranunculo crenati -Polytrichetum sexangularis (Horvat 1936) Mucina et al.nom. novum hoc loco(basionym: Polytrichetum sexangularis balcanicum Horvat 1936)

3. Soldanella pusillae-Plantaginetum gentianoidis Bo�caiu 1971(syn. Nardo-Plantaginetum gentianoidis Lakusié et al. 1979; non Nardo- Plantiaginetum gentianoidis Ganchev 1963)

4. Soldanella pusillae-Ranunculetum crenati Borza ex Bo�caiu 1971(syn.Agrostio rupestris-Ranunculteum crenati Resmerita 1975 corr. 1978,

Agrostio alpinae-Ranunculetum crenati Resmerita 1975)5. Soldanella hungaricae-Ranunculetum crenati Coldea 19856. Ranunculetum crenati Lakusié 19667. Soldanella pusillae - Luzuletum spadiceae (Borza 1934) Mucina et al. nom.

novum hoc loco(basionym: Luzuletum spadiceae retezaticum Borza 1934)

8. Soldanella hungaricae-Salicetum kitaibelianae Coldea 19859. Ligustico-Caricetum foetidae Horvat 1960 prov.

10. Thlaspio microphylli-Plantaginetum atratae Horvat 193611. Plantaginetum atratae Horvat in Horvat et al. 197 4

(syn.? Soldanello-Plantaginetum durmitorei Lakusié 1984 nom. nudum)12. Omalotheco-Alopecuretum gerardii Mucina et al. 199013. Trifolio - Phleetum pantocsekii Lakusié 1984 nom. nudum?14. Nardo - Gnaphalietum supini Bartsch 194015. Agrostio rupestris-Gnaphalietum supini Resmerita corr. 1978

(syn. Agrostio alpinae-Gnaphalietum supini Resmerità' 1975)16. Poo-Cerastietum (Soyrinki 1954) Oberdorfer 1957

Arabidetalia caeruleae Rubel 1933 (syn. Salicetalia retusae Lakusié 1968) Salicion retusae Horvat 1949

17. Saxifragetum prenjae Horvat 1931 (here?)18. Saxifrago-Rumicetum nivalis Horvat 193619. Geo-Oxyrietum digynae Horvat 193620. Bartsio-Salicetum reticulatae Mucina et al. 1990

51

21. Saxifrago semperviui-Salicetum reticulatae (Horvat 1936) Mucina et al.nom. novum hoc loco(basionym: Salicetum retuso-reticulatae macedonicum Horvat 1936)

22. Dryado-Salicetum reticulatae Beldie 1967(syn. Salicetum reticulatae sensu Pu�caru-Soroceanu 1956

23. Soldanello-Salicetum retusae Horvat 1933(syn. Soldanella - Salicetum retusae bosniacum Lakusié et al. 1979)

24. Anemono-Salicetum retusae Horvat 195325. Salicetum retuso-kitaibelianae Lakusié 197026. Gentiano-Plantaginetum atratae Mucina et al. 199027. Trifolio-Plantaginetum angustifoliae Lakusié 1966

Acknowledgements

The authors thank to ali friends who have supported us by their company during the expeditions to Bulgarian mountains, namely to P. Demes, V. Lovasova-Demesova, L'. Kost'al ,K. Husar. A. Kovér, P. Pisùt. Par! of the herbarium materia! was determined or re vis ed by J. Dvorà½_ (mainlygrasses and sedges), J. Cap (I'hymu.s), V. Grulich (Sedum), M. Kovanda (Campanula) and H. Sfposova(Galium). Some less excessible literature sources were provided by G. Caldea, R. Lakusié, Lj. Markoviéand S. Nemenz and the Library of Zoologisch -Botanische Gesellschaft in Òste1Teich. Severa! drafts ofthe paper were retyped by B. Wolfovà and later computer edited by D. Mucina, S .Mucina andB .Mucina.

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Budapest, 4 (1-2): 159-189. Stoyanov, N. & Stefanov, B. (1948) - Flora na Balgariya. Univ. Pechat., Sofiya. 1316 pp. Szafer, W .. Pawlowski, B. & Kulczyriski, S. (1927) - Die Pflanzenassoziationen des Tatra- Gebirges. III.

Teil: Die Pflanzenassoziationen des Koscieliska Tal. Bull. Acad. Polon. Sci. Lettres, Clas. Mat. Nat. B., Cracovie, II: 13-78.

Tiiuber, F. (1985) - Endemische Phytoassoziationen aus den Rumdnischen Karpathen. Fol. Geobot. Phytotax., Praha, 20 (1): 1-16.

'I\1tin, T.G.,Heywood, V .H.,Burges,N.A.,Moore,D.M., Valentine,D.H., Walters,S.M. & Webb,D.A. (edc .), (1964-1980): Flora Europaea. 1-5. Cambridge University Press, Cambridge.

Valachoviè', M. (1989) - Zur Kenntnis der Fe/sschuttvegetation an den sudlichen Hangen des PrsvivecBerges tBohinsko jezero, Jugoslawien). Acta Bot. Croat., Zagreb, 48: 103-106.

Ve!chev, V. & Vasilev, P. (1970) -Fitotsenologichno prouchvane na Kentranthus kellereri Stoj. et Stef. v Severen Pi.rin. Izv. Bot. Inst., Sofiya, 20: 123-130.

Walter, H. & Straka, H. (1970) -Arealkunde. Floristisch - historiche Geobotanik. 2. Aufl. Eugen Ulmer Verlag, Stuttgart. 478 pp.

Westhoff, V. & van der Maarel, E, (1978) -The Braun-Blanquet approach. In: Whittaker, R .H. (ed.): Class1fication of plant communities: 289-399. Dr. W. Junk, The Hague.

Wirth, V. (1980) -Flechtenflora. Eugen. Ulmer Verlag, Stuttgart. 552 pp. Yordanov, D. (ed.), (1963-1979) -Flora na NarodnaRepublikaBalgariya. I-VIII. Izd. Balg. Akad.Nauk,

Sofiya. Zollitsch, B. (1966) - Soziologische und ò'kologische Unntersuchungen auf Kalkschiefem in hocha/pinen

Gebieten. I. (Die Steinschuttgestel/schaften der Alpen). Ber. Bayer. Bot. Ges., Miichen, 40: 67-100.

The authors' addresses:

Mag. DDr. Ladislav Mu�ina, Dept. of Vegetation Ecology & Biologica! Conservation, Inst. of Plant Physiology, Vienna University, Althanstr. 14, A-1091 Wien, Austria

RNDr. Milan Valachoviè', RNDr. Ivan Jarolfmek, CSc. and RNDr. Jan Seffer, ali Dept. of Geobotany, Inst. of Experimental Biology & Ecology, Slovak Academy of Sciences, Sienkiewiczova 1, CS-814 34 Bratislava, Czcchoslovakia

RNDr. Anna Kubinska, CSc., State Centre for Natme Conservation, Dept. of Nature Conservancy Development, Heyrovského 1, CS-841 05 Bratislava, Czechoslovakia

RNDr. Ivan Pistit, CSc.,Inst. ofNatural History, Slovak National Museum, Vajanského nabrezie 2,CS-814 22 Bratislava, Czechoslovakia

56

Appendix

L ocalities of the relevés:

Tab. 1. Hieracio-Caricetum kitaibelianae (1-9) and Leontopodio-Potentilletum stojanovii (10-16).

1. Banderitsa Cottage, Djamdjievite Skali, 2000 m; 8-8-1983 (LM2694).2. Banderitsa Cottage, Djamdjievite Skali, 2050 m; 8-8-1983 (LM2695).3. Banderitsa Cottage. Djamdjievite Skali, 2050 m; 8-8-1983 (LM2696).4.Banderitsa Cottage,Djamdjievite Skali, 2100 m; 8-8-1983 (LM2697).5. Banderitsa Cottage, Djamdjievite Skali, 1900 m; 8-8-1984 (LM3107).6. Ridge between Banderitsa Cottage and Malkiya Kazan, 2100 m; 9-8-1983 (LM2699).7. Banderitsa Cottage, Djamdjievite Skali, 1900 m; 8-8-1984 (LM3106).8. Banderitsa Cottage, direction Vikhren, 1980 m; 8-8-1984 (LM3110).9. Malkiya Kazan, direction Banderitsa Cottage, 2000 m; 9-8-1984 (LM3113).

10. Golemiya Kazan, Zaslon Eltepe, 2450 m; 9-8-1984 (LM3117).11. Golemiya Kazan, flanks of Kutelo Mts., above Zaslon Eltepe, 2500 m; 10-8-1984 (LM3133).12. Saddle Kabata, 2500 m; 9-8-1983 (LM2707).13. Banderitsa Cottage, Djamdjievite Skali, 2300 m; 8-8-1983 (LM2698).14.Malkiya Kazan, 2300 m; 9-8-1983 (LM2702).15. Ridge between Malkiya and Golemiya Kazan, 2380 m; 10-8-1984 (LM3123).16. Malkiya Kazan, 2300 m; 9-8-1983 (LM2700).

Tab. 3. Silene pusilla-Saxifraga oppositifolia community.

1. Ridge between Banki Sukhodol Mt and Bayuvi Dupki Mt., 2600 m; 13-8-1984 (LM3157).2. Ridge between Bayuvi Dupki Mt. and Kamenititsa Mt., 2600 m; 13-8-1984 (LM3158).

Tab. 4. Papaveri-Armerietum alpinae (1-23) and Veronico-Silenetum prostratae (24-29).

1. Golemiya Kazan, 2450 m; 10-8-1984 (LM3127).2. Golemiya Kazan, 2420 M; 10-8-1984 (LM3132).3. Tsirkus Kamenititsa, flanks of Bayuvi Dupki Mt., 2750 m; 12-8-1984 (LM3150).4. Tsirkus Kamenititsa, saddle between Bayuvi Dupki Mt. and Kamenititsa Mts., 2650 m; 12-8-1984

(LM3156);5. Golemiya Kazan, 2500 m; 10-8-1984 (LM3130).6. Tsirkus Bayuvi Dupki, 2680 m; 11-8-1984 (LM3145).7. Golemiya Kazan, 2400 m; 10-8-1984 (LM3125).8. Tsirkus Kamenititsa, 2500 m; 12-8-1984 (LM3152).9. Tsirkus Bayuvi Dupki, 2680 m; 11-8-1984 (LM3146).

10. Tsirkus Bayuvi Dupki, 2650 m; 11-8-1984 (LM3147).11. Malkiya Kazan, 2200 m; 9-8-1984 (LM3115).12. Golemiya Kazan, 2500 m; 10-8-1984 (LM3128).13. Golemiya Kazan, S of Zaslon Eltepe, 2500 m; 10-8-1984 (LM3129).14. Golemiya Kazan, bottom, 2500 m; 10-8-1984 (LM3120).15. Golemiya Kazan, 2350 m; 10-8-1984 (LM3141).16. Malkiya Kazan, 2200 m; 9-8-1984 (LM3114).17. Tsirkus Kamenititsa, 2500 m; 12-8-1984 (LM3153).18. Golemiya Kazan, 2400 m; 10-8-1984 (LM3124).19.Kutelo Mt., 2700 m; 11-8-1984 (LM3143).20. Koncheto Ridge, 2700 m; 11-8-1984 (LM3144).21. Tsirkus Bayuvi Dupki, 2620 m; 11-8-1984 (LM3148).22. Golemiya Kazan, 2400 m; 10-8-1984 (LM3140).23. Tsirkus Kamenititsa, saddle between Bayuvi Dupki Mt. and Kamenititsa Mt., 2650 m; 13-8-1984

(LM3159).24. Golemiya Kazan, 2400 m 10-8-1984 (LM3139).25. Golemiya Kazan, 2400 m 10-8-1984 (LM3138).26. Golemiya Kazan, 2400 m 10-8-1984 (LM3137).

57

27. Golemiya Kazan, 2450 m 10-8-1984 (LM3134).28. Golemiya Kazan, 2450 m 10-8-1984 (LM3135).29. Golemiya Kazan, 2400 m 10-8-1984 (LM3136).

Tab. 5. Bromo-Geranietum macrorrhizi.

1. Banderitsa Cottage, gorge near Baikushevata mura (old specimen of Pinus leucodermi.s ), 1850 m; 8-8-1984 (LM3105).

2.Banderitsa Cottage, 1900 m; 9-8-1984 (LM3111).3.Banderitsa Cottage, direction Malkiya Kazan, 1950 m; 9-8-1984 (LM3112).4. Banderitsa Cottage, 1950 m; 8-8-1984 (LM3109).5. Banderitsa Cottage, 1930 m; 8-8-1984 (LM3108).

Tab. 6. Ligustico-Plantaginetum gentianoidis (1-4) and Omalotheco-Alopecuretum gerardii (5-21).

1. Smirnenski Tsirkus, Mitrovo Ezero; 6-8-1983 (LM2681).2. Smirnenski Tsirkus, Mitrovo Ezero; 6-8-1983 (LM2683).3. Smirnenski Tsirkus, Mitrovo Ezero; 6-8-1983 (LM2684).4. Tsirkus Belemeto, 2450 m; 7-8-1983 (LM2687).5. Saddle between Kralev Dvor Mt. and Kamenitsa Mt., 2550 m; 2-8-1978 (LM1619).7. Saddle between Kralev Dvor Mt. and Kamenitsa Mt., 2550 m; 2-8-1978 (LM1621).8. South flank of Mozgovishki Chukar Mt., 2520 m; 7-8-1983 (LM2690).9. Saddle Solishcheto (between Arabski Grab Mt. and Kuklite Mt.), 2410 m; 31-7-1978 (LM1599).

10. Saddle Solishcheto, 2410; 31-7-1978 (LM1608).11. Saddle Solishcheto, 2350 ; 31-7-1978 (LM1610).12. Tevnoto Ezero, Zaslon, 2510 m; 1-8-1978 (LM1615).13. Saddle between Kralev Dvor Mt. and Kamenitsa Mt., 2600 m; 2-8-1978 (LM 1618).14. Saddle between Kralev Dvor Mt. and Kamenitsa Mt., 2550 m; 2-8-1978 (LM1620).15. Smirnenski Tsirkus, below Kralevodvorska Porta; 7-8-1983 (LM2685).16. Tsirkus Belemeto, 2500 m; 7-8-1983 (LM2686).17. Tsirkus Belemeto, 2500 m; 7-8-1983 (LM2688).18. Treta Reka Valley; 6-8-1983 (LM2678).19. Treta Reka Valley; 6-8-1983 (LM2679).20. Smirnenski Tsirkus, Mitrovo Ezero; 6-8-1983 (LM2680).21. Ridge between Malk Tipits Mt. and Tipits Mt., 2600 m; 7-8-1983 (LM2691).

Tab. 8. Gentiano-Plantaginetum atratae (1-9) and Bartsio-Salicetum reticulatae (10-16).

1. Kamenishki Vrakh Mt. (2533 m), 2400 m; 7-8-1978 (LM 1654).2. Premkata Saddle, 2600 m; 11-8-1984 (LM3142).3. Tsirkus Kamenititsa, 2550 m; 12-8-1984 (LM3154).4. Tsirkus Kamenititsa, 2550 m; 12-8-1984 (LM3155).5. Premkata Saddle, 2610 m; 9-8-1983 (LM2706).6. Kabata Tsirkus, 2450 m; 9-8-1983 (LM2708).7. Golemiya Kazan, 2400 m; 9-8-1983 (LM2701).8. Golemiya Kazan, Zaslon Eltepe, 2400 m; 9-8-1984 (LM3118).9. Golemiya Kazan, Zaslon Eltepe, 2400 m; 9-8-1984 (LM3119).

10. Kazana Tsirkus, 2400 m; 9-8-1983 (LM2703).11. Ridge between Malkiya and Golemiya Kazan, 2350 m; 9-8-1984 (LM3116).12. Ridge between Malkiya and Golemiya Kazan, 2400 m; 10-8-1984 (LM3121).13. Ridge between Malkiya and Golemiya Kazan, 2350 m; 10-8-1984 (LM3122).14. Golemiya Kazan, 2450 m; 10-8-1984 (LM3126).15. Golemiya Kazan, 2500 m; 10-8-1984 (LM3131).16. Tsirkus Kamenititsa, flanks of Bayuvi Dupki Mt., 2750 m; 12-8-1984 (LM3149).

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the vegetation of rock fissures, screes, and … · 2.materials and methods 2.1. study area the...

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